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HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
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BREVIORA
MUSEUM OF COMPARATIVE ZOOLOGY
Harvard University
NUMBERS 351-379
1970-1971
CAMBRIDGE, MASSACHUSETTS, U.S.A.
1971
Edited
by
Penelope Lasnik
CONTENTS
BREVIORA
Museum of Comparative Zoology
Numbers 351-379
1970
No. 351. A new species in the nomeid fish genus Psenes from
the equatorial eastern Pacific. By Richard L. Haed-
rich. 7 pp. June 12.
No. 352. The Chaiiares (Argentina) Triassic reptile fauna VII,
The postcranial skeleton of the traversodontid
Massetognathus pascuali (Therapsida, Cynodontia).
By Parish A. Jenkins, Jr. 28 pp. July 10.
No. 353. Anolis jacare Boulenger, a solitary anole from the
Andes of Venezuela. By Ernest E. Williams, Osvaldo
A. Reig, Pablo Kiblisky, and Carlos Rivero-Blanco.
15 pp. August 10.
No. 354. Taxonomic and ecological notes on some Middle and
South American lizards of the genus Ameiva
(Teiidae). By Arthur C. Echternacht. 9 pp.
September 18.
No. 355. Generic relations and speciation patterns in the Cara-
caras (Aves: Falconidae). By Francois Vuilleumier.
29 pp. November 30.
No. 356. On a new species in a new earthworm genus from Puerto
Rico. By G. E. Gates. 11 pp. November 30.
No. 357. A review of the fossil Pelomedusidae (Testudines,
Pleurodira) of Asia. By Roger Conant Wood. 24
pp. November 30.
No. 358. South American anoles: Anolis apollinmis Boulenger
1919, a relative of A. biporcatus Wiegmann (Sauria,
Iguanidae). By Ernest E. Williams. 11 pp.
November 30.
No. 359. The swimbladder as a juvenile organ in stromateoid
fishes. By Michael H. Horn. 9 pp. November 30.
No. 360. Mammals from the early Cenozoic of Chubut, Argen-
tina. By George Gaylord Simpson. 1 3 pp. Novem-
ber 30.
No. 361. Additions to knowledge of the Argyrolagidae (Mam-
malia, Marsupialia) from the late Cenozoic of
Argentina. By George Gaylord Simpson. 9 pp.
November 30,
No. 362. Addition to knowledge of Groeberia (Mammalia,
Marsupialia) from the mid-Cenozoic of Argentina.
By George Gaylord Simpson. 17 pp. November 30.
1971
No. 363. Non-specificity of host-selection in the ectoparasitic
snail Odostomia (Menestho) bisuturalis (Say) (Gas-
tropoda: Pyramidellidae). By Robert C. Bullock
and Kenneth J. Boss. 7 pp. January 8.
No. 364. A new scincid lizard from Bougainville, Solomon
Islands. By Allen E. Greer and Fred Parker. 1 1 pp.
January 8.
No. 365. Characters and synonymies among the genera of ants.
Part IV. Some genera of subfamily Myrmicinae
(Hymenoptera: Formicidae). By William L. Brown,
Jr. 5 pp. January 15,
No. 366. Pulsed sound of the porpoise Lagenorhynchus australis.
By William E. Schevill and William A. Watkins. 10
pp. January 15.
No. 367. Micromischodus sugillatus, a new hemiodontid characin
fish from Brazil, and its relationship to the Chilo-
dontidae. By Tyson R. Roberts. 25 pp. January
15.
No. 368. Structural habitats of West Indian Anolis lizards I.
Lowland Jamaica. By Thomas W. and Amy
Schoener. 53 pp. January 29.
No. 369. Lithophaga aristata in the shell-plates of chitons (Mol-
lusca). By Robert C. Bullock and Kenneth J. Boss.
10 pp. January 29.
No. 370. Ecological observations on a little known South
American anole: Tropidodactyliis onca. By James
P. Collins. 6 pp. March 31.
No. 371. A new species of bromeliad-inhabiting galliwasp
(Sauria: Anguidae) from Jamaica. By Albert
Schwartz. 10 pp. March 31.
No. 372. The paleontology and evolution of Cerion II: age and
fauna of Indian shell middens on Curacao and Aruba.
By Stephen Jay Gould. 26 pp. March 31.
No. 373. The Chanares (Argentina) Triassic reptile fauna. VIII.
A fragmentary skull of a large thecodont, Liipero-
suchus fractus. By Alfred Sherwood Romer. 8 pp.
March 31.
No. 374. The fishes of the Malaysian family Phallostethidae
(Atheriniformes). By Tyson R. Roberts. 27 pp.
June 15.
No. 375. Structural habitats of West Indian Anolis lizards II.
Puerto Rican uplands. By Thomas W. and Amy
Schoener. 39 pp. June 15.
No. 376. Podocnemis venezuelensis, a new fossil pelomedusid
(Testudines, Pleurodira) from the Pliocene of
Venezuela and a review of the history of Podocnemis
in South America. By Roger Conant Wood and
Maria Lourdes Diaz de Gamero. 23 pp. June 15.
No. 377. The Chanares (Argentina) Triassic reptile fauna IX.
The Chaiiares Formation. By Alfred Sherwood
Romer. 8 pp. June 15.
No. 378. The Chanares (Argentina) Triassic reptile fauna X.
Two new but incompletely known long-limbed
pseudosuchians. By Alfred Sherwood Romer. 10
pp. June 15.
No. 379. The Chanares (Argentina) Triassic reptile fauna XL
Two new long-snouted thecodonts, Chanaresuchus
and GuaiosLichus. By Alfred Sherwood Romer. 22
pp. June 15.
INDEX OF AUTHORS
BREVIORA
Museum of Comparative Zoology
Numbers 351-379
1970-1971
No.
Boss, Kenneth J 363, 369
Brown, William, Jr 365
Bullock, Robert C 363, 369
Collins, James P 370
Diaz de Gamero, Maria Lourdes 376
Echternacht, Arthur C 354
Gates, G. E 356
Gould, Stephen Jay 372
Greer, Allen E 364
Haedrich, Richard L 351
Horn, Michael H 359
Jenkins, Parish A., Jr 352
KiBLisKY, Pablo 353
Parker, Fred 364
Reig, Osvaldo a 353
Rivero-Blanco, Carlos 353
No.
Roberts, Tyson R 367^ 374
RoMER, Alfred Sherwood 373. 377, 378, 379
ScHEviLL, William E 366
Schoener, Thomas W. and Amy 368. 375
Schwartz, Albert 37I
Simpson, George Gaylord 360, 361, 362
Vuilleumier, Francois 355
Watkins, William A 366
{j'li^^^'''' .
BREVIORA
Mmseuim of Compsirative Zoology
Cambridge, Mass. 12 June, 1970 Number 351
A NEW SPECIES IN THE NOMEID FISH GENUS PSENES
FROM THE EQUATORIAL EASTERN PACIFIC^
Richard L. Haedrich-
Abstract. Psenes sio n. sp. is based on five specimens 23-66 mm SL
from the equatorial eastern Pacific Ocean. The new species belongs to the
species-group (others are P. pelliicidiis, P. maculatus, and P. arafurensis)
with large, laterally compressed, knifelike teeth in the lower jaw. P. cya-
nophrys and P. whiteleggii have small conical teeth in both jaws. The new
species is characterised by its light color, long pelvic fins, two weak anal
spines and 23-24 rays, 18-19 pectoral finrays, and 36-38 vertebrae.
In a recent unpublished yet widely distributed manuscript
(Haedrich and Horn, 1969), a new species of Psenes was included
in a key (p. 36). It was stated explicitly that use of the name in
the key did not constitute publication, and it was indicated that a
formal description would soon appear in a review of the entire
genus. Other responsibilities, however, have virtually brought this
work to a halt, and early completion of the review does not seem
likely. Therefore, in order to avoid the nomenclatural debacle that
I see developing, this note has been prepared describing the new
species.
The specimens upon which the new species is based were made
available by J0rgen Nielsen, and are housed in Universitetets Zoo-
logiske Museum, Copenhagen (ZMC). They were collected by
Dr. Nielsen on Step-I, a cruise conducted by the Scripps Institution
of Oceanography. The manuscript has been read by Richard H.
Backus and Giles W. Mead. Figure 1 was drawn by E. Leenders.
1 Contribution No. 2486 from the Woods Hole Oceanographic Institution.
2 Woods Hole Oceanographic Institution, Woods Hole, Mass., and
Museum of Comparative Zoology, Harvard.
2 BREVIORA No. 351
Portions of this work were supported by a United States Govern-
ment Grant under the Fulbright-Hays Act, the Johs. Schmidt
Fund, and National Science Foundation grant GB- 15764.
Among stromateoid fishes, the genus Psenes is distinguished in
having two dorsal fins with the first dorsal originating over or
before the pectoral insertion, persistent thoracic pelvic fins, a deep
to moderately elongate body, and teeth present in some species on
the palatines and/or basibranchials but never on the glossohyal.
The genus, its relationships, and the nominal species were treated
in a general way by Haedrich (1967), though recent findings will
modify this account somewhat. Within Psenes, there seem to be
two species groups. One group (including P. cyanophrys and
P. whiteleggii) is characterized by small, conical, slightly recurved
teeth in both the upper and lower jaw. The other group (in-
cluding P. pellucidus, P. arajurensis, P. maciilatus and the new
species) has small, conical, slightly recurved teeth in the upper
jaw and large, laterally flattened, bladelike teeth in the lower jaw.
In recognition of the considerable contributions to marine ich-
thyology made by the Scripps Institution of Oceanography, the new
species will be known as
Psenes sio n. sp.
Figure 1
Material. Five specimens, 23-66 mm SL, all in ZMC; those
marked with an asterisk* have been X-rayed: *1 spec, 60 mm SL,
HOLOTYPE, Step-I sta. 23, 11°10'S 80°01'W, 17 Oct. 1960,
2250-2345 hrs., 0-90 m, 5' net, surf. temp. 17.8°C. PARA-
TYPES: *1 spec, 66 mm SL, Step-I sta. 80-1, 1°24'S 94°55'W,
2/3 Dec. 1960, 2335-0230 hrs., high-speed net, battered. 2 spec,
26 & 44 mm SL, Step-I sta. 73-1, 4°22'S 95°04'W, 2 Dec. 1960,
0200-0500 hrs., high-speed net, very battered, smaller spec,
cleaned-and-stained. *1 spec, 23 mm SL, Step-I sta. 80, 1°59'S
94°55'W, 2 Dec. 1960, 2200 hrs., dipnet-nightlight, surf. temp.
22°C. All specimens are immature.
Diagnosis. An elongate, compressed, light-colored Psenes with
large, compressed close-set teeth in the lower jaw, long pelvic fins,
two weak anal spines and 23-24 rays, and 36 to 38 vertebrae.
Description. Individual proportions and counts are presented in
Table 1.
The body is elongate, the maximum depth of larger specimens
1970 NEW NOMEID FISH 3
being around 30 per cent of the standard length, and is very com-
pressed. The caudal peduncle is tapered, compressed, and some-
what elongate. The musculature, though firm, is translucent, par-
ticularly along the anal fin base and over the viscera. The two
dorsal fins are scarcely divided. The first dorsal fin originates over
the edge of the opercle and comprises 10 to 12 thin brittle spines,
the second originates just behind mid-body and comprises 23 to 25
long rays. The entire fin folds partially into a very shallow groove
and terminates behind the end of the anal fin. The anus is at mid-
body, in a slit. The anal fin commences shortly behind the anus
under the third or fourth ray of the second dorsal, and is composed
of two weak spines and 23 or 24 long rays. The muscles for ele-
vating the anal rays can be clearly seen, as can the basal elements
of both median fins. The pectoral fin is long and fairly broad, with
18 or 19 rays; its base is incUned about 45° to the vertical. The
pelvic fins are very long; they insert under the middle or end of
the pectoral fin base and extend beyond the anal origin, and are
composed of one short spine and five long branched rays. The
caudal fin, broken in most specimens, is apparently long and
forked; the small elements preceding the principal rays extend
well forward on the peduncle. The cycloid scales are very small,
and do not appear to extend significantly onto the bases of the
median fins. The scales are extremely deciduous, and most are
gone; the count of scale pockets along the lateral line of the holo-
type is ca. 85. The skin is thin; the subdermal mucus canal system
is but httle developed, and the body pores are very small or
wanting.
The head is around 35 per cent of the standard length, its profile
sloping. The skin of the top of the head is naked, and pores are
clearly visible, particularly those over the head of the hyomandibu-
lar. The eye is of moderate size, located a little more than its
diameter from the tip of the truncate snout, and does not enter
into the profile of the head. There is apparently no adipose tissue
around the eye, but somes does extend forward from the front of
the eye to surround the nostrils. The two small nostrils are located
much nearer to the tip of the snout than to the eye. The end of
the maxillary is below the anterior border of the eye, but the angle
of the gape is well before the eye. The premaxillary is not pro-
tractile. The lacrimal bone is large and transparent, and covers
the top of the maxillary. The teeth are uniserial in the jaws. The
teeth in the upper jaw are small, conical, slightly recurved, and
spaced; the teeth in the lower jaw are large, at least twice as long
4 BREVioRA No. 351
as those in the upper jaw, compressed and knifeUke with very small
cusps, and very close-set. The vomer, palatines, and basibranchials
appear to be toothless. The oral valves are prominent. The oper-
cles are very thin; their margins are either entire or set with ex-
tremely fine spinules. The striated opercle has two very weak flat
spines; the angle of the preopercle is rounded but does not bulge
backward. The gill-rakers are moderate, blunt, about half the
length of the filaments, and bear fine teeth on their inner edges; the
rakers are spaced, about 15 on the lower limb of the first arch.
The pseudobranch is well developed, but there are no rudimentary
rakers below it. The light yellow thymus is clearly visible. There
are six branchiostegal rays.
The color in alcohol is tan, darker on the back than on the sides.
There is a suggestion of three brownish vertical bands on the after
part of the body in some specimens. The first dorsal fin is dark,
but all the other fins are whitish. The dark lining of the gill cavity
shows clearly through the transparent opercles. The dark peri-
toneum shows clearly through the thin abdominal wall. The in-
side of the mouth is fight yellow, and the eye is grey.
The skeleton in general is very light. This is particularly ap-
parent in the thin transparent dermal skeleton. There is no supra-
maxillary bone. The supraoccipital is but little developed. There
is a wide opening between the cleithrum and the coracoid. The
pelvic bones reach to the cleithrum. There is a large foramen in
the scapula. The postcleithrum can be plainly seen through the
body wall, and it extends to the lower margin of the body. There
are 36 to 38 vertebrae, including the hypural; about 12 to 15 verte-
brae appear to be precaudal. In the tail, there are two autogenous
haemal spines, four hypurals, two paired uroneurals, and three
epurals. The second and third hypurals are broad triangular plates
much larger than any other caudal element. The first hypural bears
an hypuropophysis. Three free interneurals precede the dorsal fin.
The first dorsal interneural supports two spines. The two anal
spines are weak.
Remarks. Most species of P series are widely distributed in the
tropical and sub-tropical parts of the world ocean. P. sio, how-
ever, is quite restricted, and has been found only in the eastern
Pacific from about 11° S (holotype) to perhaps 10° N (R. Rosen-
blatt, additional Scripps specimens, in lift.). P. sio most closely
resembles P. rnaculatus, an apparently antitropical species known
only from sub-tropical waters in the North and South Atlantic.
Within the range of Psenes sio, only P. cyanophrys is known to
occur for sure. This species may be distinguished from P. sio
1970 NEW NOMEID FISH 5
by its small conical teeth that are similar in both jaws (as men-
tioned above), its color pattern of fine horizontal stripes (P. sio
has about three indistinct vertical bands), its greater maximum
depth (43-52% SL vs. 29-41% SL in P. sio), its generally greater
number of median fin rays (D 24-29 vs. 23-25, A 24-28 vs.
23-24), and its fewer vertebrae (31 v^. 36-38).
As yet unrecorded from the eastern tropical Pacific but known
from the western parts of that ocean are Psenes pellucidus and
P. arafurensis. The former has more median finrays and verte-
brae (D 27-32, A 26-31, vert. 41-42) than P. sio, the latter fewer
(D 18-22, A 20-22, vert. 31).
Psenes whiteleggii, from the Indian Ocean and Australia, has
conical teeth in both jaws and low median finray and vertebral
counts (D 17-20, A 17-18, vert. 31-32). The Atlantic P. macu-
latus, the species most similar to P. sio, has slightly fewer median
finrays and vertebrae (D 22-24, A 22-24, vert. 35) and more
pectoral finrays (21-22 vs. 19 in P. sio) and anal spines (III vs.
II). The preanal distance is 58-63% SL in P. maculatus, and
51-54% SL in P. sio.
LITERATURE CITED
Haedrich, Richard L. 1967. The stromateoid fishes: systematics and a
classification. Bull. Mus. Comp. Zool., Harvard, 135(2): 31-139.
Haedrich, Richard L., and Michael H. Horn. 1969. A key to the
stromateoid fishes. Woods Hole Oceanographic Institution Ref. No.
69-70, September 1969, 46 pp. Unpublished Manuscript.
(Received 2 April 1970.)
6 BREVIORA No. 351
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BREVIORA
Miiseimi of Cointipairsitive Zoology
Cambridge. Mass. 10 July, 1970 Number 352
THE CHANARES (ARGENTINA) TRIAS5IC REPTILE FAUNA
VII. THE P05TCRANIAL SKELETON OF THE TRAVERSO-
DONTID MASSETOGNATHUS PASCUALI (THERAPSIDA,
CYNODONTIA)
Parish A. Jenkins, Jr.^
Abstract. The postcranial skeleton of Massetognathus pascuali is
described from a single nearly complete individual and four disarticulated
specimens; manus, pes and pubis could not be described from the available
material. M. pascuali has a postcranial skeleton morphologically similar to
that in Pascualgnathns polanskii (an earlier traversodontid) and in African
Triassic cynodonts. The basic skeletal pattern of all these forms differs
from that in Exaeretodon sp., an advanced traversodontid. The specialized,
imbricating ribs of M. pascuali are intermediate in form between those
of P. polanskii (which are diademodontid in form) and those of Exaere-
todon sp. (which essentially have lost the cynodont specialization).
Although Richard Owen made the first study of a cynodont more
than a century ago, a detailed knowledge of cynodonts has accumu-
lated only recently. First known only from Africa, representatives
of this advanced group of mammallike reptiles have now been
recovered from Asia and North and South America. The most
specialized and, during the early and middle Triassic, the most
abundant cynodonts were the herbivorous gomphodonts. Trans-
versely broad, multicuspid postcanine molars with heavy occlusal
wear (often to the point of effacing the original crown pattern)
are characteristic of the group. Romer (1967) recognized three
gomphodont families: Diademodontidae, Traversodontidae, and
Trirhachodontidae. Massetognathus pascuali is a traversodontid
and is one of two species described by Romer (1967) from the
' Department of Anatomy, Columbia University, New York, New
York 10032
2 BREVIORA No. 352
Chanares Formation of Argentina. Other undoubted traversodon-
tid genera include Traversodon from the Santa Maria Formation of
Brasil, Exaeretodon, Proexaeretodon and Ischignathus from the
Ischigualasto Formation of Argentina, and Scalenodon and Scal-
enodontoides from the Manda and Molteno beds, respectively, of
Africa. Pasciialgnathus polanskii from the Puesto Viejo Forma-
tion of Argentina, originally classified as a diademodontid (Bona-
parte, 1966a; 1966b), is now interpreted as a primitive traverso-
dontid on the basis of tooth morphology (Bonaparte, 1967, and
personal communication).
The present account of the Massetognathus pasciiali postcranial
skeleton is based upon the disarticulated remains of at least four
individuals and an almost completely articulated skeleton of a fifth
individual, all about the same size and preserved in a nodule several
inches thick and two and one-half feet in diameter. The 1964-65
expedition of the Museo de la Plata and Museum of Comparative
Zoology collected the nodule from the Chanares Formation in the
Chanares-Gualo region of western Argentina. All the skeletal
material is catalogued as No. 3691 in the Museum of Compara-
tive Zoology.
Although the available postcranial material of Massetognathus
pasciiali is incomplete, there are two reasons why even a partial
account is important. First, some workers regard cynodonts as
ancestral to mammals (Crompton and Jenkins, 1968; Hopson and
Crompton, 1969). Gomphodont cynodonts are certainly not di-
rectly related to mammals, but they undoubtedly possessed a de-
gree of biological organization that at least approximated that of
their carnivorous-insectivorous relatives (from some form of which
mammals were derived). Thus all cynodonts are relevant to evalu-
ating the reptile-mammal transition. Second, most published in-
vestigations of cynodonts have been restricted to cranial anatomy.
Well-preserved, generically determinate postcranial skeletons are
rare. Of South American genera, only Belesodon (von Huene,
1935-1942), Exaeretodon (Bonaparte, 1963a) and Pasciialgna-
thus are known from relatively complete skeletons; the preservation
of known Belesodon material is poor, however, and Exaeretodon
appears to be morphologically aberrant in comparison to the gen-
eral pattern known from other cynodonts. Postcranial material
associated with Traversodon and Chiniquodon (von Huene, 1935
1942) is only fragmentary, although Romer (1969) has recently
described more complete limb bones of Chiniquodon. I have re-
viewed the postcranial skeletons of African cynodonts (Jenkins, in
1970
MASSETOGNATHUS POSTCRANIAL SKELETON
press). Of the Russian forms very little is known; some data is
available for Permocyiiodon (Konjukova, 1946).
Vertebral column and ribs
The number of presacral vertebrae in Massetognathus pascuali
is at least 23 and is here interpreted to be about 26. A hiatus in
the lumbar series of the one nearly complete, articulated vertebral
column is responsible for this uncertainty. Present are 7 cervicals,
164-? dorsals, and 6 sacrals. Judging from the length of several
articulated but incomplete caudal series, tail vertebrae numbered
at least 17 and probably as many as 25. A count of 26 presacral
vertebrae in M. pascuali compares with 26 in Leavachia duven-
hagei (Broom, 1948), 27 in Thrinaxodon liorhinus (Jenkins, in
press), 28 in Exaeretodon sp. (Bonaparte, 1963a), 29 in Cynog-
nathus crateronotus (Seeley, 1895 ) and at least 30 in a large cyno-
dont identified tentatively as Diademodon sp. (Brink, 1955; Jen-
kins, in press).
Two separately articulating atlas arches appear to conform to
the general pattern known from African Triassic cynodonts (Jen-
kins, in press), but their preservation is too poor to allow specific
description or comparison. The atlas intercentrum. transversely
elongate and ventrally convex, bears a posteriorly directed process
or lip (1, Fig. IB). A concave articular facet on the dorsal sur-
face of this lip contacts a median convex facet on the anterior
aspect of the atlas centrum (f a i, Fig. 6B ) . The lateral extremities
of the intercentrum each bear, on their dorsal surfaces, a concave
facet that articulates with the ventral aspect of one of the two
occipital condyles. The atlas centrum (6 mm long) is synostosed
Figure 1. Vertebral elements in Massetognathus pascuali. A. Cervical
vertebrae, probably the third and fourth, in lateral view. B. First or atlantal
intercentrum in ventral view with anterior margin toward the top of the
page. C, Anterior dorsal ("thoracic") vertebrae; vertebra on the right is
in lateral view, on the left posterolateral view. All X 1. Abbreviations:
fo, costal fovea for rib head; 1. posterior lip of the intercentrum; p. para-
pophysis; t, transverse process.
4 BREVIORA No. 352
to the axis centrum (8.5 mm long), forming a conspicuously large
vertebra. On the dorso-lateral aspects of the atlas centrum are two
facets, one for each atlas arch half (f a a. Fig. 6B). Protruding
from the median aspect of the centrum between these facets is a
small process (d, Fig. 6B) homologous with the dens or odontoid
process of mammals. Elsewhere (Jenkins, 1969; in press) I have
proposed that the mammalian dens originated as a neomorphic
process from the atlas centrum of cynodonts. The occurrence of a
dens in Massetognathus pascuali is unequivocal and supports the
theory that both a dens and an atlas centrum (of which the dens
was formerly thought to be a vestige) occur together among
cynodonts.
The axis centrum of the one articulated cervical series is about
9 mm long, 1 mm longer than the centra of the following cervicals.
The axial prezygapophyses have been broken off on all specimens,
but from the appearance of their narrow bases^ — no thicker than
the laminae from which they protrude — they are vestigial. Axial
postzygapophyses of normal size bear articular facets at an angle
estimated to be 25° to the horizontal. The axial spine, a flat blade
with a distinctly mammalian shape, exhibits a straight dorsal mar-
gin that was probably slightly convex during hfe. The robust
transverse processes are directed posterolaterally.
Cervical centra are amphicoelous, as are all other centra in
Massetognathus pascuali, with the exception of the atlas (only the
posterior aspect bears a notochordal concavity) and possibly some
of the smaller caudal vertebrae. In cervical vertebrae, the rim sur-
rounding the notochordal concavity on each end is swollen, and
this increases the concave curvature of the sides and ventral aspect
of each centrum. A bulbous parapophysis is to be found on the
ventrolateral aspect of each anterior rim (p. Fig. lA). If the rib
head articulated on the apex of the parapophysis, as seems most
likely on the basis of the large size of the process, then this condi-
tion differs from that in species of Thrinaxodon and Cynognathus,
in which cervical rib heads are situated intervertebrally. A median
ventral keel, similar to that in other cynodont cervicals, traverses
the length of each cervical centrum.
Cervical pedicles are narrow anteroposteriorly and bear short,
stout transverse processes (t, Fig. lA). Cervical spines, broken
off on all but two disarticulated cervicals (Fig. lA), are trans-
versely slender and recurved, and taper abruptly toward the apex.
Of particular interest are the zygapophyses, which provide a basis
for distinction between cervical and dorsal (anterior thoracic)
vertebrae. As noted above, the axial postzygapophyseal facets are
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 5
oriented at an angle of about 25° to the horizontal; the orientation
of posterior facets on the third through sixth cervical are more than
25° — probably about 35°. The distance between their lateral mar-
gins is approximately 7 mm. The anterior articular facets of the
seventh cervical of course conform in orientation and spacing to
those of the foregoing series. The posterior articular facets, how-
ever, appear to be oriented at about 45°. Those on the succeeding
(eighth) vertebra are nearly parasagittal — i.e., within a few de-
grees of vertical — and are only 3.5 mm apart. The articular facet
characteristics of the eighth vertebra, continued (with gradual
modification) through the dorsal series, definitely estabhsh a dif-
ferent pattern. Although transitional, the seventh vertebra most
closely resembles other neck vertebrae and thus may be regarded
as the last cervical. Similar changes in facet orientation and
spacing, together with other morphological changes, occur between
the seventh and eighth vertebrae of Tlvinaxodon liorhimis and
Cynognathiis craterouotiis (Jenkins, in press). The condition in
Massetognathus pascuali reaffirms the fact that the "mammalian"
number of seven cervical vertebrae was already established in
cynodonts.
With the exception of the atlas intercentrum, no other cervical
intercentra have been identified. The broad grooves formed by the
rims of adjacent centra are evidence that intercentra were present —
as in Thrinaxodon liorhimis, for example — and the state of dis-
articulation accounts for their postmortem loss.
The dorsal vertebral column in cynodonts is either a relatively
undifferentiated dorsal series (as in most reptiles) or two series —
"thoracic" and "lumbar" (as in mammals). In certain African
Triassic cynodonts, separation of thoracic and lumbar regions may
be made on the basis of distinct morphological specializations of
the posterior dorsal (= lumbar) ribs (Jenkins, in press). In other
cynodonts (e.g., Exaeretodon sp.; Bonaparte, 1963a) the trunk
cannot be divided into thoracic and lumbar regions because the
posterior ribs are not specialized. Massetognathus pascuali has
specialized "lumbar" ribs and clearly belongs to the first category.
The exact number of thoracic and lumbar vertebrae is unknown
because a complete vertebral column with ribs is not yet available.
Therefore, thoracic and lumbar vertebrae will be given only a
general description as anterior and posterior dorsals, respectively,
although the ribs (to be discussed below) clearly give evidence of
a differentiated series.
The centra of anterior dorsal vertebrae are approximately 8.5
mm long; those of posterior dorsals are about 1 1 mm long (10 mm
6 BREVIORA No. 352
in the one articulated specimen). The costal foveae (fo. Fig. IC)
of anterior dorsal vertebrae form a cleft for reception of the rib
head, which is therefore intervertebral in position. On posterior
dorsals the rib head articulates with a parapophysis (p, Fig. 2B)
and is not intervertebral. Transverse processes are laminar and
bowed dorsally on anterior dorsal vertebrae (t, Fig. IC) but are
rodlike and round in cross-section on posterior dorsals (t, Fig. 2B).
The posterior intervertebral notch is deep throughout the dorsal
series, the anterior notch negligible or absent. In contrast to many
cynodonts, anapophyses are lacking. Dorsal prezygapophyses are
robust, extending to or slightly beyond the level of the anterior
aspect of the centrum. Articular facets on the first dorsal vertebra
are about 3.5 mm apart and on the ninth are 4.0 mm; an abrupt
widening takes place at some point in the middle or posterior dorsal
series, for the articular facets of the penultimate dorsal vertebra
are about 7 mm apart. The narrowly-spaced facets of anterior
dorsals are nearly vertical, whereas those wider apart on the pos-
terior dorsals are oriented at angles of as much as 45°. Neural
spines on anterior dorsals are narrow, but unlike cervical spines,
do not taper significantly at the apex; they incline caudad at angles
of about 30°. Spines on posterior dorsal vertebrae are broad an-
teroposteriorly, leaving only a narrow gap between vertebrae; their
inclination is only a few degrees caudad.
Sacral vertebrae successively decrease in size posteriorly. The
centrum of the first sacral is approximately as long as those of the
posterior dorsals (about 10 mm), while the sixth and last sacral
centrum is some 2 mm shorter. Massive synapophyses — repre-
senting fused parapophyses and transverse processes — arise from
the pedicles and from the anterior half of the side of the centrum.
The zygapophyses, proportionately less robust than in the dorsal
series, diminish in size on successively more posterior sacrals. In
contrast to the orientation and spacing in the posterior dorsals,
sacral articular facets incline nearly parasagittally and are nar-
rowly spaced. At the last dorsal-first sacral articulation, the facets
are 5.5 mm apart, but between the third and fourth and fourth and
fifth sacrals they are only about 2 mm apart. At the fifth-sixth
sacral articulation the trend is reversed; the facets are spaced
3.5 mm apart and appear to be inclined at 10° from the vertical.
The trend toward less verticality of facets is continued into the
caudal series. Sacral spines successively diminish in height and
anteroposterior breadth. The spine on the first sacral is approxi-
mately 8 mm in height and 7 mm in breadth (versus 8.5 mm height
and 8 mm breadth for the last dorsal spine). On the fifth sacral
i
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 7
the same measurements are 5.5 mm and 4.2 mm respectively. This
trend is continued into the caudal series. The apices of sacral
spines tend to be oval in contrast to those of the posterior dorsals
which are elongate and attentuated at each end.
Caudal centra decrease in length from 6.5 mm at the first caudal
to about 5 mm at the fifth. More posterior caudals preserved with
MCZ 3691 are disarticulated, and their position cannot be posi-
tively assigned. However, one isolated series of eleven caudals
shows a decrease in centrum length from 5 mm (which is evidence
that it is approximately the fifth caudal) at the first to 4 mm at
the last (? fifteenth caudal). Other specimens show that at least the
first five caudals bear synapophyses. Articular facets are inclined
at approximately 45°, at least through the first five caudals. The
width between the lateral edges of these facets decreases from
6 mm (between the last sacral and first caudal) to 4.5 mm
(between the fourth and fifth caudals). The terminal caudal is
unknown.
Massetognathus pascuali probably possessed ribs on all presacral
vertebrae, as in other cynodonts for which adequate material is
known. Ribs were not found in association with the first six cervi-
cal vertebrae, although the morphology of the transverse processes
and parapophyses on the axis through sixth cervical is clear evi-
dence of their existence. These features cannot be verified at pres-
ent on the atlas. The ribs of the seventh cervical appear to have
been shorter than, but otherwise similar to, those of the anterior
dorsal series.
Dorsal ribs in Massetognathus pascuali are of basically two
types: in the anterior and middle dorsal series, a freely articulating
rib of normal costal form and proportions, and in the posterior
dorsal series, a fused rib with a Y-shaped distal end. Unfortu-
nately, the transitional ribs between the two types are as yet
unknown.
The proximal ends of anterior and middle dorsal ribs are basi-
cally triangular. The tuberculum and capitulum form two corners
of the triangle; the shaft arises from the third (Fig. 3A). On the
anterior aspect of this triangular surface is a slight crest (c. Fig.
3A) comparable to a similar feature on the anterior dorsal ribs of
Cynognathus sp. (c. Fig. 3B. C). There is no other apparent fea-
ture that makes this type of rib distinctive. The maximum widths of
the shafts near their proximal ends are between 2.5 and 3 mm. The
distal ends, as preserved, are approximately 1.5 mm thick. Ribs of
this basic morphology (as opposed to the specialized posterior
8
BREVIORA
No. 352
Figure 2. The posterior dorsal and sacral region in Massetoguathus
pasciiali in A, dorsal and B, lateral views. X 1. Abbreviations: a il,
acetabular facet of the ilium; a p, anterior process of rib shaft; b, ridge
on dorsal aspect of rib shaft; f a p, articular facet on anterior process of
rib shaft; gr, groove on dorsal margin of ischium; il, ilium; is, ischium;
is t, ischial tuberosity; m p, medial process on base of ilium for articulation
with ischium and pubis; p, parapophysis; p p, posterior process of rib shaft;
t, transverse process.
dorsal ribs to be described next) are associated with at least the
first thirteen dorsal vertebrae.
The following description of the specialized posterior dorsal
ribs is based on the penultimate and last dorsal ribs preserved in
articulation (Fig. 2) and in addition, a few disarticulated pieces of
similar morphology. These ribs have a short shaft that bifurcates
into a Y-shaped terminus with two processes (ap, pp, Figs. 2, 4B).
The anterior process is broader than the posterior and bears, on
the dorsal surface of its tip, a rather flat, round facet (f a p, Figs.
2A, 4B). Articulating with this facet is the end of the posterior
process of the preceding rib. Presumably, the underside of the
970
MASSETOGNATHUS POSTCRANIAL SKELETON
Figure 3. Proximal ends of dorsal ribs of A, Masseto^natliiis pascuali
(X 2), and of B and C, Cynognatlms sp. (X V2) in anterior view. Ab-
breviations: c, crest on anterior aspect of shaft (see text for details).
posterior process also bears a facet. The rib as a whole projects
laterad from the vertebral column and appears not to have had any
ventral curvature. In lateral view (Fig. 2B) the shaft and point of
bifurcation are at approximately the same level, but the anterior
and posterior processes incline somewhat ventrally. A low, bony
ridge (b. Figs. 2 A, 4B) runs obliquely onto the posterior process
from the point of bifurcation where it is most prominent. This
feature is comparable to a similar ridge on the ribs of Cynognatlms
sp. (b. Fig. 4A) and other cynodonts; in the fourteenth thoracic
Figure 4. Specialized posterior dorsal ("lumbar") ribs of various
cynodonts. A. Cynognathus crateronotiis (British Museum of Natural
History no. R. 2571), X V4. B, Massetogucithiis pascuali. XL C,
Leavacliia duvenhagei (Rubidge Collection, Graaf Reinet, South Africa,
no. 92), X V2. All dorsal views. Abbreviations: a p, anterior process
of rib; b, ridge on dorsal aspect of rib shaft (of unknown function but
probably homologous in the forms shown here); f a p, articular facet on
anterior process of rib shaft; LI, L2, first and second lumbar vertebrae;
p p, posterior process of rib shaft; T14, T15, fourteenth and fifteenth
thoracic vertebrae.
10 BREVIORA No. 352
rib of C (T14, Fig. 4A), the ridge is merely a linear elevation on
the flat costal plate. On successive ribs, however, the ridge becomes
more prominent until, in the lumbar ribs (LI, L2, Fig. 4A), it
reflects forward to contact the preceding rib plate (see Jenkins, in
press). No such reflection is evident in Massetognathus pascuali,
but the ridge morphology and general pattern of the process are
nevertheless similar to that of the fourteenth and fifteenth thoracic
rib plates of C
Specialized, imbricating ribs are common but not universal
among cynodonts. Known members of the earliest cynodont fam-
ily, the procynosuchids, apparently did not possess this specializa-
tion (see discussion below, however, for a possible exception).
Galesaurids, typified by the well known Thrinaxodon liorhinus
(Jenkins, in press), developed costal expansions on all presacral
ribs. Members of three other families, e.g., Cynogmitlms craterono-
tus (Cynognathidae; Seeley, 1895), Diadeinodon sp. (Diademo-
dontidae; Jenkins, in press) and Cricodon metabolus (Triracho-
dontidae; Crompton, 1955), possessed imbricating ribs only in the
posterior dorsal region. The ribs in chiniquodontids (von Huene,
1935-1942) are as yet unknown. There remains only the Traver-
sodontidae, which Bonaparte (1963b) characterized, on the basis
of species of Exaeretodon and supposedly Traversodon, as lacking
synostosed ribs with overlapping processes. For this and other
reasons, Bonaparte interpreted traversodontids as probably having
arisen from procynosuchids along a lineage separate from that an-
cestral to all other cynodont families (whose members possess rib
specializations). However, Massetognathus pascuali unquestion-
ably possesses synostosed lumbar ribs with details comparable to
the Cynognathus-Diademodon pattern. Pascualgnathus polanskii,
now classified as a traversodontid (Bonaparte, 1967), has lumbar
ribs that are unquestionably diademodontid in pattern. Further-
more, von Huene (1935-1942; 137-140) described expanded ribs
("Facherrippen") synostosed to the lumbar vertebrae in Traverso-
don stahleckeri. Crompton (1955) presented circumstantial evi-
dence that the traversodont Scalenodon from the African Manda
beds also possessed the expanded rib specialization. Yet Bona-
parte (1963a) amply demonstrated that at least one traversodont,
Exaeretodon sp., did not possess such specialization. Presacral ribs
in Exaeretodon sp. are morphologically uniform and are more or
less freely articulating (although the more "solid" attachment of
the last three dorsal ribs, as described by Bonaparte, possibly rep-
resents a vestige of a less mobile articulation typical of expanded
ribs ) . In view of this unexpected association at the family level of
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 11
forms possessing and forms lacking rib specializations, the taxo-
nomic significance of this character should be reassessed. Further
comment is reserved for the discussion below.
Sacral ribs in Massetognathiis pascuali have an essentially con-
fluent capitulum and tuberculum, a short shaft and an expanded
distal end for articulation with the iliac blade. Proximally each rib
is synostosed to its corresponding vertebra. If I may judge from
the disarticulated condition of every known sacroiliac joint, liga-
ments and cartilage must have been chiefly responsible for binding
the ilium and sacral ribs. The concave distal ends of the sacral ribs
conform to the gently convex internal surface of the ilium, but
they do not appear to form any osseous interdigitation by which
sacroiliac joints are commonly reinforced. Viewed from above, the
distal end of the first sacral rib (Si, left side. Fig. 2A) is Y-shaped
with processes directed anterolaterad and posterolaterad. Articu-
lating with the dorsal surface of the anterolateral process is the
posterior process of the last dorsal (lumbar) rib. The second sacral
rib has the largest distal expansion; irregularly shaped and widest
anteriorly, the expansion is 7 mm long in one well preserved speci-
men. The third, fourth and fifth ribs bear more or less symmetrical
distal expansions that are successively smaller caudally. The first
four sacral rib shafts are oriented more or less laterally, the fifth
slightly anterolaterally. The sixth and last sacral rib bears a shaft
that is directed posterolaterally and a bifurcated, Y-shaped termi-
nus resembling that of posterior dorsal ribs. Its iliac articular sur-
face is narrow and strap shaped; the rib and vertebra could well be
interpreted as the first caudal were it not for its position opposite
the posterior tip of the ihac blade (S6, Fig. 2A). While it is appar-
ent that the width between the distal ends of this pair of ribs
(20 mm) is much less than that of the fourth and fifth sacrals
(27 mm), the intervening gap could well have been completed by
ligaments. This interpretation of the sixth sacral vertebra and ribs
may be open to future modification, but on present evidence ap-
pears to be the most probable.
Only the first three caudal ribs are preserved. All are synostosed,
their shafts directly posterolaterally. The first is only 8 mm long
(Cdl, Fig. 2A), with two blunt processes on its terminus resembling
a stunted version of the last sacral rib. As far as can be deter-
mined, terminal processes were not developed on the second and
third caudal ribs, which are 5 mm or less in length. Ribs on suc-
ceeding vertebrae must have been very small and probably did not
occur in the posterior caudal series.
12
BREVIORA
No. 352
Shoulder Girdle
Available interclavicles of Massetognatlius pascuali are incom-
plete, although there is sufficient material to conclude that the
morphology is very similar to that in Thrinaxodon liorhinus and
different from that in Exaeretodon sp. In outline the interclavicle
is cruciate with an elongate posterior ramus (pr, Fig. 5A). The
entire bone, although basically a flat plate, is bowed ventrally from
front to back. Two ridges, one longitudinal, the other transverse,
divide the ventral surface of the interclavicle into quadrants. The
two anterior quadrants (c c. Fig. 5A) are shallow concavities for
reception of the proximal ends of the clavicles. The ridges are
#^f
prox
Figure 5. A, The interclavicle and B, the right clavicle of Massetog-
nathus pascuali, both in ventral view. X 1. Abbreviations: ca, concavity
for acromion; c c, concavity for proximal end of the clavicle; dist, distal
end of clavicle; f, ventral flange on distal end of clavicle; pr. posterior
ramus of interclavicle; prox, proximal end of clavicle.
most prominent at their intersection. With the exception of the
posterior part of the longitudinal ridge, which gradually fades out,
the ridges become more salient toward the margins. There is no
evidence that the longitudinal ridge was a deep keel as in Exaereto-
don sp. (Bonaparte, 1963a). The posterior ramus in M. pascuali
is similar in length and form to that in galesaurids, and is unlike the
very short ramus of Exaeretodon sp.
The Massetognathus pascuali clavicle is robust. The proximal
two-thirds are more or less straight (Fig. 5B), the distal third
curving sharply posterodorsally. The broad, flat plate on the proxi-
mal end articulates with the previously described concavity (c c.
Fig. 5A) in the interclavicle. Along the ventral aspect of the distal
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 13
third runs a flange (f, Fig. 5B) similar to that noted in African
Triassic cynodonts (Jenkins, in press) and Exaeretodon sp. (Bona-
parte, 1963a). This flange continues to the distal end where it
contributes to the formation of a concavity (ca, Fig. 5B) for re-
ception of the acromion. The clavicle is essentially identical to
that in galesaurids.
The scapula, coracoid, and procoracoid in Massetognathiis pas-
ciicili are firmly synostosed, although the joints can readily be dis-
tinguished (Fig. 6A). The scapular blade, elongate and narrow,
bears a distinct concavity on its lateral surface — a fossa presumably
for the supracoracoideus muscle, the infraspinatus homologue of
mammals. The anterior margin of the blade is reflected sharply
laterally, the posterior margin somewhat less so. An acromion
process as such is not preserved on any of the scapulae; this ab-
sence may be due to postmortem damage to a delicate process or
to the fact that the clavicular concavity simply fitted to the convex
edge of the anterior scapular base (ac, Fig. 6A). The scapular
half of the glenoid is a hemicircular and shghtly convex facet that
faces posteroventrally and somewhat laterally.
The coracoid is basically triangular in lateral view (co. Fig. 6A).
The posterior end forms an elongate, attenuated process terminated
by a tubercle for the origin of the coracoid head of the triceps (co
tr, Fig. 6A ) . The process is morphologically similar to the same
feature in Pascualgnathus pokmskii and in African Triassic cyno-
donts, but differs in form from that in Exaeretodon sp. as inter-
preted by Bonaparte ( 1963a). Between the glenoid and the triceps
tubercle, the superior margin of the coracoid is about 2 mm wide
and is slightly concave from front to back. The slightly convex
inferior margin is, in contrast, extremely thin bone, and, as a con-
sequence, is invariably damaged postmortem. A saddle-shaped
facet, concave dorsoventrally and convex transversely, constitutes
the coracoid half of the glenoid.
The procoracoid (pr, Fig. 6A) appears to be an irregularly
shaped, flat plate, but in no available specimen are the free mar-
gins complete. There are so many basic similarities in the scapu-
locoracoid of Massetognathiis pascuali and African cynodonts that
a complete procoracoid of the former would probably have the
same oval shape characteristic of the latter. A crescentic depres-
sion on the lateral aspect along the inferior margin may represent
the biceps origin. Above lies the round procoracoid foramen
(f pr. Fig. 6A) sculptured in a ventrolateral direction to facilitate
passage of its nerve and blood vessels. The evidence as to
whether the procoracoid participated in the glenoid is equivocal. I
14
BREVIORA
No. 352
believe that the procoracoid probably supported articular cartilage
at the very anterior extremity of the glenoid (see left glenoid,
Fig. 6A), but there is no certainty of the procoracoid contribu-
ting to the shoulder joint as there is for some African Triassic
cvnodonts.
970 MASSETOGNATHUS POSTCRANIAL SKELETON 15
Figure 6. Elements of the postcranial skeleton of Massetogiuithiis
pasciiali, drawn as preserved in situ. A, Incomplete left shoulder girdle and
forelimbs seen from the left side. B. Lateral view of axis. C, Dorsal view
of left humerus. D, Medial view of left radius and ulna. All X 1. Ab-
breviations: ac, area of clavicular articulation (distinct acromion process
not developed); cl. clavicle; co, coracoid; co tr, tubercle for coracoid head
of triceps; cp. capitellum; d, dens; d f. distal flange on radius; dp, delto-
pectoral flange; f a a, atlas centrum facet for atlas arch; f a i, atlas
centrum facet for atlas intercentrum; f ec, ectepicondylar foramen; f en,
entepicondylar foramen; f pr, procoracoid foramen; g, groove possibly
representing teres major insertion or the origin of one of the humeral
triceps heads; h, humeral head; hu, humerus; 1, ridge possibly representing
insertion of the teres minor; 1 t, lesser tuberosity; p f, proximal flange on
radius; pr. procoracoid; ra, radius; s, scapula; ul, ulna; ul f, ulnar flange.
16 BREVIORA No. 352
Forelimb
Principal characteristics of the humerus of Massetognathus pas-
ciuili are the relatively broad expansion of the proximal and distal
ends, and the large deltopectoral flange. The proximal end of the
shaft is bowed dorsally and the head is oriented to a more dorsal
position. The well-rounded head (h, Fig. 6C) possesses greatest
curvature along its dorsoventral axis. As preserved, the articular
surface is confluent medially whh the lesser tuberosity (1 t, Fig.
6C) and laterally with the proximal margin of the deltopectoral
flange. The greater tuberosity presumably arose in the mammalian
lineage between the head and the proximal margin of the delto-
pectoral crest, but in M. pasciiali there is no evidence of a distinct
tubercle. The greatest width of the proximal end, from the lesser
tuberosity to the region of the presumptive greater tuberosity,
measures approximately one-third the total length of the humerus.
The broad deltopectoral flange is slightly more than half the total
length of the humerus. The free edge of the flange thickens and
everts (laterally) at its proximal and distal extremities, but along
the middle part is rather thin and flat. From the region of the pre-
sumptive greater tuberosity a low, bony ridge runs obliquely across
the flange toward the shaft (1, Fig. 6C). An identical ridge on the
humeri of certain African Triassic cynodonts has been interpreted
as possibly representing the insertion of a teres minor (Jenkins, in
press). On the posterodorsal aspect of the shaft is a groove pos-
sibly representing the insertion of the teres major or the origin of
one of the humeral triceps heads (g. Fig. 6C); Bonaparte ( 1966b)
interpreted a rugosity at this site in Pascualgnathus polcmskii as the
origin of the medial triceps head.
The distal end of the humerus is triangular in dorsal view, its
maximum breadth being approximately 40 per cent of the humeral
length. Arising from the robust ectepicondylar region, a thin supra-
condylar flange runs proximally as well as somewhat dorsally. The
flange, pierced in its proximal half by a small ectepicondylar fora-
men (f ec, Fig. 6A, C), becomes a low crest at the middle of the
shaft and is continuous with the ridge (1, Fig. 6C) described above.
A stout bar of bone arising from the entepicondylar region encloses
an elongate, oval entepicondylar foramen (f en. Fig. 6A, C). The
capitellum (cp, Fig. 6A) is bulbous and contributes to the thick-
ness of the ectepicondylar region. The trochlea immediately
adjacent is a broad, shallow groove; the principal axis of this
groove is dorsoventral, as expected, but it is also slightly
oblique — the dorsal part being more laterally situated than the
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 17
ventral part. Morphologically, the humerus of M. pasciudi is essen-
tially identical to that in Pasciuil^naihus polcmskii and galesaurids;
apparent differences with galesaurids, e.g., the greater roundness
of the head and capitellum, are due to the better ossification in
M. pascuali and P. polcmskii. As Bonaparte (1963a) noted, the
humerus of Exaeretodon sp. is more similar to the dicynodont
or gorgonopsid pattern than to that typical of galesaurids, and
thus stands in contrast to the conventional cynodont humerus of
M. pascuali.
The radius has a slight sigmoidal curvature (which facilitates its
crossing over the anterior aspect of the ulna) and expanded proxi-
mal and distal ends. The nearly circular proximal articular facet
forms a shallow concavity, in which the greatest curvature is an-
teroposterior (as is its reciprocal surface on the capitellum). On
the posteromedial aspect of the proximal end is an excrescence that
bears a facet (f u. Fig. 6D) apparently for articulation with the
ulna. From this excrescence, a distinct flange (p f. Fig. 6D) runs
distally to about the midpoint of the shaft. On better preserved
material of African Triassic cynodonts, I interpreted a similar
flange as possibly being associated with the biceps insertion and
the radio-ulnar interosseous ligament (Jenkins, in press). Bona-
parte (1963a) interpreted a similar feature in Exaeretodon sp. as
marking the position of the interosseous ligament. The distal end
of the radius expands gradually to the distal articular facet which
is oval (long axis transverse) and shallowly concave. A distal
flange ( d f. Fig. 6D ) arises near the midpoint of the shaft essen-
tially as a continuation of the attenuating proximal flange described
above. Beginning on the posterior aspect of the shaft, the distal
flange takes a spiral course toward the lateral aspect as it enlarges
distally. Its position is suggestive of the attachment of an inter-
muscular septum separating flexor and extensor muscle groups.
The ulna, like the radius, is sigmoidally shaped but is expanded
only at its proximal end (Fig. 6D). The relatively shallow semi-
lunar notch represents the typical cynodont condition — basically
oval in outline, but with a rather straight medial margin and a
nearly hemicircular lateral margin. An olecranon process is not
present or at least was not ossified; the proximal end of the ulna,
where such a process would be developed, is broad and rugose.
The transversely narrow shaft of the ulna bears on its anteromedial
aspect a flange (ul f. Fig. 6D) that extends from the semilunar
notch to the distal articular facet. In all probability this flange rep-
resents the ulnar attachment of the interosseous ligament. The
lateral surface of the shaft reveals one large, spoon-shaped fossa
proximally, and on the medial surface two fossae, one proximal,
the other distal. All three fossae are wefl represented in African
18 BREVIORA No. 352
Triassic cynodonts, and I have proposed (Jenkins, in press) that
they represent origins of various manual flexor and extensor mus-
cles. The distal articular facet, convex from front to back, is broad
anteriorly and narrow posteriorly and thus is triangular in outline.
Both the radius and ulna of M. pascuali, as far as available mate-
rial permits comparison, are extremely similar to their counterparts
among galesaurids and in Pascualgnathus polanskii. Although
definite similarities exist with the antebrachial elements of Exaere-
todon sp., the essential identity of the M. /7<;/.vcwa//-galesaurid pat-
tern is incontrovertible.
Only an incomplete and disarticulated series of seven or eight
carpals of Massetognathus pascuali is known, but these are so
poorly preserved, and good comparative material is so scanty, that
no constructive observations on the manus can be made at this
time.
Pelvis
A complete pelvis of Massetognathus pascuali is not yet avail-
able, although enough is known of the ilium and ischium for pre-
liminary description and comparison with other forms. The ilium
bears an elongate, vertical blade, spatulate in front and lanceolate
behind (il. Fig. 2). The lateral aspect of the blade is concave,
especially anteriorly. The shape of blade, as well as the relative
proportions of the pre- and postacetabular regions, is most similar
to that in Pascualgnathus polanskii and is comparable to that of
galesaurids, cynognathids, and diademodontids; Exaeretodon sp.,
on the other hand, has an iliac blade quite unlike the foregoing
(Bonaparte, 1963a). The base of the iliac blade in M. pascuali is
constricted into a short neck, below which are medial and lateral
processes. The medial process (m p. Fig. 2B) bears two articular
surfaces — one each for the pubis and ischium — which intersect at
an angle of about 150°. The lateral process bears a nearly circular,
concave facet that represents the iliac contribution to the acetabu-
lum (a il. Fig. 2B). The facet is oriented principally in a postero-
ventral direction but with a slight lateral eversion.
Only the dorsal half of the ischium is available for examination
(is. Fig. 2). The concave acetabular surface faces anterolaterally
and is oriented essentially vertically. The postacetabular part of the
ischium constitutes a broad plate that ventrally meets its counter-
part of the opposite side. A longitudinal groove (gr. Fig. 2) on
the dorsal margin of this plate terminates posteriorly at an ischial
tuberosity (is t. Fig. 2).
Available pubes have been extensively damaged postmortem.
970
MASSETOGNATHUS POSTCRANIAL SKELETON
19
The pubic contribution to the acetabulum is considerably smaller
than that of the ischium. Neither the obturator fenestra nor the
ventral aspect of the pelvic basin is preserved.
The bony acetabulum is relatively shallow — a little more than
5 mm deep. The continuous, sharp rim around the acetabulum de-
fines a more circular socket than that known in African Triassic
genera. In all other details, the pelvis of Massetognathus pasciiali
appears to be morphologically similar to that in Pascualgnathus
polanskii, galesaurids, and even larger African Triassic forms.
Hindlimb
The femur (Figs. 7, 8A) is a moderately slender bone, except
for the expanded proximal end, which bears robust trochanters.
The femoral head, bulbous and almost hemispherical as in mam-
mals, is reflected medially but also somewhat dorsally by virtue of
the dorsal bowing of the proximal end of the shaft (Fig. 8 A ) . The
protuberant trochanter major (tr mj, Figs. 7, 8A) measures about
5 mm in thickness. A pear-shaped area of smooth bone on its apex
may represent the principal site of muscle attachment or of a sub-
tendinous bursa. The bone surface immediately adjacent to the
apex is rugose. The trochanter minor (tr mn. Figs. 7, 8A) forms
an elongate flange that arises abruptly near the intertrochanteric
tr mj
tr mn tr mn
tr mj
Figure 7. Reconstruction of a left femur in Massetognathus pasciiali
in A, ventral, and B, dorsal views. X 1. Abbreviations: i f. intertrochanteric
fossa; tr mj. trochanter major: tr mn. trochanter minor.
20
BREVIORA
No. 352
fossa (i f. Fig. 7) and gradually disappears slightly distal to the
shaft's midpoint. In cross-section, the middle of the shaft is essen-
tially rectangular; its thickness from extensor (dorsal) to flexor
(ventral) surfaces is about 6 mm, from the medial to lateral sur-
faces 4.5 mm. The distal end of the femur expands gradually but
asymmetrically, the lateral condyle being broader and farther offset
from the femoral axis than the medial. The medial condyle pro-
jects more ventrally than does the lateral condyle. The fibula ap-
pears to have articulated on the lateral epicondylar region where a
shallow groove (f f, Fig. 8A) occurs. The femur of Massetog-
nathus pascuoli is morphologically comparable to the femora of
Pascualgnathus pokmskii and even the larger African genera in
which ossification of the extremities was well developed (Jenkins,
in press). In smaller forms, such as galesaurids, the femur appears
to be diiTerent because of the lack of ossification of the extremities
and trochanters.
The tibia is a transversely slender bone, bowed somewhat an-
teriorly. The two proximal articular facets, oval in outline and
shallowly concave, are separated by a low, median ridge. The
B
Figure 8. Elements of the hindlimb of Massetognathiis pascuoli. drawn
as preserved in situ. A. Right femur, tibia and fibula in lateral view. B.
Left tibia in medial view. All X 1. Abbreviations: f f. facet for articula-
tion with fibula; fl, fibular flange for femoral articulation; f t, lateral tibial
fossa of uncertain significance; g t, groove on medial aspect of tibia; 1 t.
lineation on lateral aspect of tibia; tr mj, trochanter major; tr mn.
trochanter minor.
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 21
lateral margin of the lateral facet is thickened and protuberant, and
it appears likely that part of the proximal fibula articulated here.
On the proximal end of the lateral aspect of the shaft is a deep
fossa (f t, Fig. 8A) of uncertain significance; from the posterior
margin of this fossa a faint lineation (1 t. Fig. 8A) runs obliquely
across the shaft to merge with the narrow anterior margin of the
shaft. Also of uncertain significance is a slightly curved groove
(g t. Fig. 8B) along the middle of the shaft's medial aspect. The
distal extremity of the tibia has a marked lateral expansion that
sufficiently widens at the terminus to accommodate a broad,
slightly convex facet for the astragalus.
The fibula, a very slender-shafted bone with enlarged extremi-
ties, is bowed laterally. The shaft appears to have been wider
transversely than anteroposteriorly. On the one complete speci-
men, a shallow groove running the length of the shaft along its
medial aspect can be detected. A flange on the proximal end
(fl. Fig. 8A) probably contacted a groove on the lateral epicon-
dylar region of the femur; the remainder of the proximal fibula
articulated with the tibia. The distal end, like the proximal, broad-
ens anteroposteriorly and is inflected somewhat medially. All
features of both tibia and fibula found in MassetognatJius pascuali
are duplicated in galesaurids and even in the larger African Triassic
genera. The tibia and fibula of Exaeretodon sp., by contrast, de-
part from the uniform pattern of other cynodonts by being pro-
portionally more massive.
Nothing can be described of the pes of Massetognathus pascuali
from the available material.
Discussion
Massetognathus pascuali, about 50 cm in length from head to
tail, was a cynodont of relatively slender build (Fig. 9). The short-
ness of the limbs relative to the approximated trunk length gives
the body a "low-slung" appearance. The head seems dispropor-
tionately large for the body, but a relatively massive head is a
common cynodont characteristic.
The postcranial skeleton of Massetognathus pascuali is basically
like that in galesaurids, diademodontids, cynognathids and Pas-
cualgnathus polanskii. Limited morphological diversity appears to
be the rule for the postcranial skeletons of Triassic cynodonts.
Exaeretodon, however, is an exception; the postcranial skeleton in
this genus differs in major details from the pattern characteristic
of other Triassic cynodonts. Bonaparte (1963a) recognized post-
cranial specializations in his original description and implied that
22
BREVIORA
No. 352
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1970 MASSETOGNATHUS POSTCRANIAL SKELETON 23
similar features might also be characteristic of other traversodon-
tids. From similarities between the genera Exaeretodon and Lea-
vachia, Bonaparte further inferred that traversodontids possibly
evolved from procynosuchids in a lineage separate from that giving
rise to other cynodont families. These conclusions, credible on
evidence available in 1963, now require modification, because
P. polanskii and M. pasciiali demonstrate that Exaeretodon is a
specialized traversodontid. Such specialization is not surprising in
view of the fact that this genus lived later in the Triassic than any
form with which it has been compared. Limb bones in Exaereto-
don sp. are more robust than in other cynodonts (Fig. 10), no
doubt in relation to its unusually large size. Some Exaeretodon
features are unknown in other cynodonts. For example, a deep,
sagittal keel occurs on an unusually short interclavicle; an extra
trochanteric process runs distally from the trochanter major along
the ventral aspect of the femur (Bonaparte, 1963a; a similar
process, identified as a trochanter minor by Parrington (1961), is
known in a whaitsid therocephalian and in a scaloposaurid bauri-
amorph). Furthermore, the lack of rib specialization is a unique
feature for a Triassic cynodont. Thus, as a specialized traverso-
dontid, Exaeretodon is no longer as relevant to the problem of
traversodontid origins as it was when other traversodontid genera
were poorly known.
Two facts bear on the problem of traversodontid origins. First,
morphological similarities between the postcranial skeletons of
Massetognathus pasciiali, Pasciialgnathiis polanskii, and African
Triassic cynodonts are evidence of close phylogenetic relationship.
Similarities between the skull and specialized ribs of P. polanskii
and Diadeinodon (see Bonaparte 1966b) raise the likelihood that
traversodontids and diademodontids were derived from a common
stock. Second, the paucity of information available on the pro-
cynosuchid postcranial skeleton does not appear to be useful in
evaluating relationships with other cynodont families. Bonaparte
( 1963a) compared the postcranial morphology of Exaeretodon sp.
and Leavachia duvenhagei and on this basis suggested the possibility
that traversodontids and procynosuchids were directly related. The
difficulty of this approach may be illustrated with respect to the
forelimb in which, for example, Bonaparte noted similarity between
the radius and ulna of Exaeretodon sp. and L. duvenhagei. The
radius and ulna of E. duvenhagei are, in fact, poorly known and in
relative proportions do not resemble those in Exaeretodon sp. more
than those of African Triassic cynodonts (Fig. 10). Similarities
cited by Bonaparte between the manus of Exaeretodon sp. and
24
BREVIORA
No. 352
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 25
Figure 10. Diagrammatic comparison of postcranial bones in various
major groups of cynodonts. Leavachia, in part modified from Broom
(1948) and in part drawn from stereoscopic photographs, represents
Procynosuchidae. The galesaurid and cynognathid-diademodontid pat-
terns are taken from Jenkins (in press); cynognathids and diademodontids
are sufficiently alike in their postcranial morphology to be represented
here by a single pattern. The Exaeretodon material is drawn from Bona-
parte (1963b). All bones have been reduced to a standard dimension to
illustrate proportional differences.
26 BREVIORA No. 352
L. diivenhagei are of equivocal significance in view of the almost
total lack of information on the manus of other cynodonts. Bona-
parte's contention that the humeri of Exaeretodon sp. and L. dii-
venhagei are proportioned alike is valid, but the dissimilarity to
those in other cynodonts is not so great as to make this comparison
especially significant (Fig. 10). Bonaparte admitted that the
scapulocoracoid of Exaeretodon sp. is more like that in Cynog-
nathus sp. than that in L. diivenhagei. This resemblance, in Fig-
ure 10 at least, is not particularly marked, ahhough the speciahzed
character of the scapulocoracoid in Exaeretodon sp. is evident.
Thus, comparisons between forelimb features of several cynodont
families provide no evidence of special affinity between traverso-
dontids and procynosuchids.
The iliac blade of Massetognathus pascuali has a long posterior
process like that in African Triassic cynodonts (Fig. 10) and
Pascualgnathiis polanskii. In contrast, the same process in Exaere-
todon sp. is relatively short (Fig. 10), and as a consequence the
blade as a whole has a more mammahan shape (Bonaparte,
1963a). The iliac blade in the type of Leavachia diivenhagei ap-
pears to have a short posterior process (Fig. 10; Broom, 1948) but
also appears to be somewhat damaged. Thus a long posterior
process may have existed in procynosuchids, and on present evi-
dence, at least, the ilia of Exaeretodon sp. and L. duvenhagei do
not indicate special affinity between the two.
Bonaparte (1963a) cited the absence of specialized ribs in
Exaeretodon sp. and Leavachia duvenhagei as possible evidence of
a close relationship between traversodontids and procynosuchids.
The subsequent discovery of specialized ribs in Massetognathus
pascuali and the reclassification of Pascualgnathiis polanskii (which
has specialized ribs of the Diademodon pattern) make this observa-
tion no longer significant. It is interesting, however, that in the
type of Leavachia duvenhagei (Rubidge Collection No. 92, Graaf
Reinet, South Africa) the last dorsal rib appears to be laterally
oriented (Fig. 4C), much as in M. pascuali and in other cynodonts
with specialized ribs. On the dorsal aspect of the shaft is a ridge
(b, Fig. 4C) comparable in position and orientation to a similar
ridge in M. pascuali, cynognathids. and diademodontids (b. Fig.
4A, B). Furthermore. Konjukova (1946) figured a specimen of
the procynosuchid Permocynodon, incompletely prepared from the
ventral aspect. The posterior dorsal ribs are directed anterolaterad
(as in Thrinaxodon, for example) and the shafts appear to be
wider than those of more anterior dorsal ribs. The capitular articu-
lations of posterior dorsal ribs in Permocynodon are relatively
broad, a feature typical of cynodonts in which the posterior dorsal
1970 MASSETOGNATHUS POSTCRANIAL SKELETON 27
ribs are synostosed. On this incomplete evidence, it appears pos-
sible that rib specialization was already underway in procyno-
suchids and was further modified in the various cynodont families
descended from them.
Hopson and Crompton (1969), in a discussion of the origin of
mammals, observe that galesaurids such as Thrinaxodon liorhinus
would be ideal candidates for mammalian ancestors were it not for
the presence of specialized ribs. These authors suggest that Exaere-
todon is relevant to this problem because its nonspecialized ribs
may represent a reversion from a specialized pattern, and similar
trends may have occurred in galesaurids. There is now substantial
evidence favoring the view that rib specialization was gradually lost
in traversodontids. P. polanskii, the earliest traversodontid for
which ribs are known, has a costal morphology of a Diadeinodon
pattern. M. pascuali, temporally intermediate between P. polanskii
and Exaeretodon sp., has ribs with less extensive specialization.
Thus the ribs of Exaeretodon sp. apparently represent a reversion
to a nonspecialized condition, and are not primitively nonspecial-
ized as originally suggested by Bonaparte (1963a). That such a
loss occurred in one family of cynodonts increases the possibility
that a similar reversion occurred in advanced galesaurids — or their
descendants — during the reptile-mammal transition.
ACKNOWLEDGMENTS
I am grateful to Professor Alfred S. Romer of Harvard Univer-
sity for enabling me to participate in the Chanares faunal studies.
Professor Romer, together with Professor Rosendo Pascual, Uni-
versidad de la Plata (Argentina), generously consented to my re-
quest to describe Massetognathus pascuali as a sequel to my work
on African cynodonts. Professor Romer furthermore provided
preparational facilities and staff, skillfully supervised by Mr. Arnold
D. Lewis, with the result that the material was presented to me in
excellent condition. Dr. Jose F. Bonaparte of the Institute Miguel
Lillo de Tucuman (Argentina) kindly read the manuscript and
clarified several important points of cynodont anatomy and rela-
tionships. Figures 6 and 8 were prepared by Mr. Robert J.
Demarest. I thank Dr. James A. Hopson of the University of
Chicago for making available his collection of stereoscopic photo-
graphs of Traversodon, and Dr. James Kitching of the Bernard
Price Institute for Palaeontological Research (Johannesburg) for
verifying some features on a specimen of Leavachia.
The collection of the Chanares material was aided by National
Science Foundation Grant GB-2454; preparation and publication
of the results has been supported by grants GB-46 1 5 and GB-8 171.
28 BREVIORA No. 352
LITERATURE CITED
Bonaparte, J. F. 1963a. Descripcion del esqueleto postcraneano de
Exaeretodon. Acta Geol. Lilloana, 4: 5-52.
1963b. La familia Traversodontidae. Acta Geol. Lilloana,
4: 163-194.
1966a. Sobre nuevos terapsidos Triasicos hallados en el
centro de la Provincia de Mendoza, Argentina. Acta Geol. Lilloana.
8: 91-100.
1966b. Una niieva "fauna" Triasica de Argentina (The-
rapsida: Cynodontia, Dicyncdontia) consideraciones filogeneticas y
paleobiogeograficas. Ameghiniana, 4: 243-296.
1967. Los tetrapodos Triasicos de Argentina. First Inter-
national Symposium on Gondwana Stratigraphy and Paleontology,
Mar del Plata.
Brink, A. S. 1955. A study on the skeleton of Diculenwdon. Palaeont.
Afr.. 3: 3-39.
Broom, R. 1948. A contribution to our knowledge of the vertebrates
of the Karroo Beds of South Africa. Trans. Roy. Soc. Edinburgh,
61: 577-629.
Crompton, a. W. 1955. On some Triassic cynodonts from Tanganyika.
Proc. Zool. Soc. London. 125: 617-669.
Crompton. A. W., and F. A. Jenkins, Jr. 1968. Molar occlusion in
Late Triassic mammals. Biol. Rev., 43: 427-458.
HoPSON, J. A., AND A. W. Crompton. 1969. Origin of mammals. In
T. Dobzhansky, et ai. (eds. ), Evolutionary Biology, Vol. 111. New
York: Appleton-Century-Crofts, pp. 15-72.
HuENE, F. VON. 1935-1942. Die fossilien Reptilien der siidamerikanischen
Gondwanalandes. Munich, C. H. Beck'sche Verlagsbuchhandlung,
332 pp.
Jenkins, F. A. Jr. 1969. The evolution and development of the dens of
the mammalian axis. Anat. Rec, 164: 173-184.
In press. The postcranial anatomy of African cynodonts
and problems in the early evolution of the mammalian postcranial
skeleton. Bull. Peabody Mus. Nat. Hist., Yale.
KoNJUKOVA, E. D. 1946. New data on Perniocynodon siishkiiii Woodw.,
a cynodont member of the Northern Dvina fauna. Dokl. Akad. Nauk,
54: 527-530.
Parrington, F. R. 1961. The evolution of the mammalian femur. Proc.
Zool. Soc. London. 137: 285-298.
RoMER, A. S. 1967. The Chanares (Argentina) Triassic reptile fauna.
III. Two new gomphodonts, Massctogiuttluis pasciiali and M. teniggii.
Breviora, No. 264: 1-25.
1969. The Brazilian Triassic cynodont reptiles Belesodon
and Chiniqiiodoii. Breviora, No. 332: 1-16.
Seeley, H. G. 1895. Researches on the structure, organization and
classification of fossil Reptilia. Part IX, Section 5. On the skeleton in
new Cynodontia from the Karroo rocks. Phil. Trans. R. Soc, Ser. B,
186: 59-148.
(Received 8 January 1970.)
BREVIORA
leseiiiini of Comtipsirative Zoology
Cambridge, Mass. 10 August, 1970 Number 353
Anolis iacare Boulenger, a "solitcry" anole from the Andes
of Venezuela
Ernest E. Williams,^
Osvaldo A. Reig,!^-
Pablo Kiblisky,- and
Carlos Rivero-Blanco'^
Abstract. Anolis jacare Boulenger is the sole member of its genus in
the Andes of Merida in Venezuela. In external morphology, size, and to
some extent in behavior, it resembles its congeners on the one anole islands
of the Lesser Antilles. The karyotype of A. jacare. however, demonstrates
that it is not closely related to either of the two Lesser Antillean stocks
which it resembles and these we know not to be closely related to each
other. The similarity of A. jacare to the two Lesser Antillean stocks and
of these to each other seems to be due to selection for a similar ecological
type.
In 1903 Boulenger described Anolis jacare from several speci-
mens in a collection made by S. Briceno at Merida, Venezuela,
at an elevation of 1 600 meters. As all too frequently happens in
Boulenger's work, the description was altogether without com-
parison or note on relationship.
Since its description additional specimens have been taken, all
in the Venezuelan Andes, but there has been little discussion of the
species. There has never been any question of its validity.
1 Museum of Comparative Zoology. Harvard University, Cambridge, Mass.
02138
2 Institute de Zoologia Tropical, Universidad Central de Venezuela, Aptdo.
59058. Caracas. Venezuela
■= Jardin Zoologico "El Pinar." Cota 905, Caracas, Venezuela
BREVIORA
No. 353
Schmidt (1939: 9) mentioned a peculiar feature of the spscies,
the double row of keeled scales forming the dorsal caudal margin.
This is a feature which A. jacare shares with some South American
species and with the very distantly related A. harkeri of Mexico.
In 1960 Etheridge placed jacare in the latijrons series of his alpha
section of the genus Anolis. This section, distinguished by the
absence of transverse processes on the caudal vertebrae, represents
an old endemic South American stock, which today shares South
America with more recent (beta section) invaders from Central
America.
Figure 1. Anolis jacare. Dorsal view of head. AMNH 13444.
No previous mention of A. jacare has cited its most interesting
feature: alone of native South American species, it shows a
close resemblance to West Indian species, specifically those of the
Lesser Antilles. In fact, its resemblance to Anolis leachii, except
in size, is such that even an experienced student of Anolis, pre-
sented with a specimen of jacare without locality, is very likely to
confuse it with A. leachii.
Table 1 compares A. jacare to A. leachii and to the Leeward
Island species more similar in adult size. A. marmoratiis.
When A . jacare is closely examined, of course, there should be
no real possibility of confusing it with A. leachii. The dorsal
squamation of the tail, larger dewlap, relatively larger ventrals.
smaller scales on the snout, etc., permit easy recognition of v4.
1970 ANOLIS JACARE 3
jacare. On general appearance, however, relationship will still
seem plausible.
However, in terms of geography, close relationship of jacare of
the Andes of Venezuela and leachii of the northern Lesser Antilles
is prima facie unlikely. In addition, there is good evidence that
the biniaculatus group (Gorman and Atkins. 1969) is derived
from the still more distant Puerto Rican stock and that it is to this
Puerto Rican stock or a still more primitive group that any phy-
letically meaningful resemblance would be expected.
Geographically, the roquet group in the southern Lesser Antilles
would be a little more plausible as close relatives of jacare than the
biniaculatus group. Gorman and Atkins (1969) have commented
on the close external resemblance of the roquet and biniaculatus
groups. Earlier, Underwood (1959) was able to find only the
most trivial scale differences between the two groups. However,
on all the characters by which Underwood was able to separate
the roquet and bimaculatus groups, jacare fits the bimaculatus
group.
With the eight Venezuelan species geographically closest, jacare
shows little similarity. (None of the eight appear to overlap the
distribution of jacare at all.) Five of the eight {chrysolepis,
auratus, juscoauratus, tropidogaster, biporcatus) belong to the
beta division of Anolis (Etheridge, 1960) and are neither osteo-
logically nor in squamation close to jacare. The three remaining
species {squamulatus, punctatus, tigrinus) are referred by
Etheridge (1960) to the same group as jacare (the latifrons series
of the alpha section of Anolis). However, these again show no
evidence of close relationship. Squamulatus and tigrinus are very
different from jacare in size {squamulatus a giant, tigrinus a
dwarf), and punctatus is conspicuously specialized in the swollen
snout of the male. All differ significantly from jacare in scale
characters.
There are indeed no South American or other continental
snecies to which A. jacare shows important resemblances. We are
left, therefore, with the external similarities to the bimacidatus
species group and, less marked, to the roquet species group. If
these resemblances go deeper, we appear to be faced with a zoo-
geographic puzzle which may need a difficult and complex solution.
It has seemed worthwhile, therefore, to broaden the study of
A. jacare to include such more recently utilized characters as
karyotype and ecology. The remainder of this paper deals with
the results of these analyses.
4 BREVIORA No. 353
Chromososome analysis (O. A. Reig and P. Kiblisky):
Four male and three female individuals have been worked for
chromosome analysis. Our report is based on the four male
individuals. We failed to get results with one of the females, and
the other two were sent to Dr. George Gorman, who, by the use
of a blood culture microtechnique, obtained a chromosome count
agreeing with our results ( personal communication ) . The male
specimens have been deposited in the Collection of Herpetology
of the Museum of Natural History of Caracas (MCNC 5601-
5604). Those studied by Dr. George Gorman are in the Museum
of Vertebrate Zoology, University of California.
Our animals were injected with 0.5 cc Colchicine Merck (solu-
tion 5 mg per cc ) 2-3 hours before killing. Testes were re-
moved, minced with scalpel, and pretreated for 20 minutes in a
hypotonic solution of sodium citrate 0.7%. The material was
centrifuged at 800 rpm and the pellet resuspended in 3/1 alcohol-
acetic fixative. After a new centrifugation, the pellet was changed
to 2/1 fixative. Spreads were obtained by air-drying on chilled
slides or by squashing, then stained with acetolactic orcein.
Giemsa and Feulgen, and mounted in Canadian balsam. Chromo-
somes were observed with a Wild M-20 microscope, and each
appropriate metaphase or meiotic prophase was recorded and
sketched. Numerous additional cells were also counted and ob-
served. Selected cells from those recorded were photographed
with high contrast Copy Kodak film, and karyotypes were con-
structed from enlarged prints. A total of 50 cells was recorded,
as listed below:
Specimen Nr. MCNC 5601
Specimen Nr. MCNC 5602
Specimen Nr. MCNC 5603
Specimen Nr. MCNC 5604
All specimens 41 8 1
Results
The diploid chromosome complement of Anolis jacare is com-
posed of 32 chromosomes (Figs. 3 and 4). Of them, 12 are
macrochromosomes and 20 are microchromosomes. The diaki-
neses (Fig. 5 ) show six large bivalents and ten very small bivalents.
Chromosome number and structure are identical in all the studied
specimens. The six pairs of macrochromosomes do not gradually
ipermalos^onicd
i)U'l<iplia\cs
D
akineses
Metaphases II
9
3
4
1
1
15
1
13
3
1970 ANOLIS JACARE 5
decrease in size but can be divided into tliree groups (Fig. I).
Group A is formed by three pairs of large metacentric and sub-
metacentric chromosomes. (In the following, we use the nomen-
clature proposed by Levan, Fregda, and Sandberg, 1964.) Pair
A-1 comprises sin chromosomes, whereas pairs A-2 and A-3 are
//^-chromosomes. Chromosomes of pair B-1 are around 4/5 the
length of those of pair A-3. Pairs B-1 and B-2 are easily
distinguishable in size. Group C comprises one pair of small si
chromosomes (r = 3.66), clearly smaller than those of pair B-2
and three times larger than the largest chromosome of the set
of the microchromosomes. A small difference in size and arm
ratio was found in the chromosomes of this pair in all the cells
where the shape of these chromosomes was clear enough, so that
the pair might tentatively be considered as heteromorphic.
Whether or not the presumptive heteromorphic pair is to be
interpreted as an X-Y sexual system cannot be solved in the
absence of good female metaphases. In two of the chromosome
spreads obtained by Gorman, the female karyotype also shows
heteromorphism in this pair. Moreover, the ring-shaped form of
the corresponding bivalent in male diakinesis does not seem to fit
with the X-Y hypothesis. The 20 pairs of microchromosomes
steadily decrease in size and seem to have terminal (/) or sub-
terminal (st) centromeres.
Gorman (1965), Gorman and Atkins (1967, 1968a) and
Gorman, Atkins and Holzinger (1967) have demonstrated that
a karyotype of six pairs of macrochromosomes and twelve pairs
of microchromosomes is shared by most of the studied species of
iguanid lizards, including the anoles of the alpha group of
Etheridge ( 1960) other than those of the bimaculatus series. The
anoles of the beta group of Etheridge depart from this "standard"
iguanid karyotype in showing seven pairs of macrochromosomes
and a variable number of microchromosomes. Within the alpha
group, the species of the bimaculatus series known in chromosome
constitution {bimaculatus, leachii, gingivinus, and marmoratus)
(see Gorman, 1965; Gorman and Atkins, 1966) are peculiar in
having quite another kind of karyotype. In these species there is
no sharp distinction between macro- and microchromosomes.
There are from 18 to 20 chromosomes gradually decreasing in
size that continue in five or six pairs of dotlike microchromosomes.
Anolis jacare departs from both the beta anoles and alpha
anoles of the bimaculatus series in retaining the "standard" set of
six pairs of macrochromosomes, easily distinguishable from the
6 BREVIORA No. 353
microchromosome set. The morphological similarities referred
to above with members of the bimaculatus series are thus not
supported by chromosome evidence, but this evidence agrees with
the osteological evidence in indicating that this species belongs to
the alpha group. Within the non-biinaculatus series alpha anoles
so far known in chromosome structure, however, a considerable
variation occurs in details of structure of the macrochromosome
set and in the number of microchromosomes. Anolis roquet,
equestris, carolinensis, and cybotes are different from Anolis jacare
in showing 22 or 24 microchromosomes and a steady decrease in
size of the macrochromosomes, the only distinguishable break in
size in these being between the fifth and the sixth pairs. Anolis
cooki, pulchellus, cristatellus and scriptus of the cristatellus series
(Gorman, Thomas, and Atkins, 1968) show the two sharp breaks
in the macrochromosomes that are also observed in A. jacare, but
in them the second break falls between the fourth and the fifth
pairs instead of between the fifth and the sixth pairs as in A. jacare.
In addition, those species of the cristatellus series mentioned above
have heteromorphic sex chromosomes and only from 15 to 18
microchromosomes. A. trinitatis and A. aeneus of the primitive
latijrons series agree with A. jacare in the two size discontinuities
among the macrochromosomes. They have, however, 24 and 22
microchromosomes respectively, and the first break in the macro-
chromosomes falls between the second and the third pair. More-
over, the first pair of macrochromosomes is metacentric in all the
illustrated karyotypes of alpha anoles, whereas it is submetacentric
in A. jacare.
Anolis jacare thus seems to be an isolated species within the
alpha group on the basis of the pattern of the size discontinuities
among the macrochromosomes and the unique number of 20
microchromosomes. It is suggestive that a distinction of three
groups within the macrochromosomes falling in the same order as
in A. jacare can also be observed in the species of the beta anoles
of the grahami and chrysolepis series so far reported (Gorman,
1965; Gorman and Atkins, 1967). There is, however, a sharp
difference between the macrochromosome set of these species and
that of A. jacare: in the former the group C comprises two pairs
instead of one pair as in the latter, the number of pairs of macro-
chromosomes thus amounting to a total of seven, as in all of the
beta anoles.
1970 ANOLIS JACARE 7
Given the widespread occurrence of six pairs of macrochromo-
somes in alpha anoles and most iguanids, we are inclined to evalu-
ate differences in number of the macrochromosome set as more
important than structural rearrangements within this portion of the
karyotype. For this reason, and because A. jcicare is clearly an
aipha Anolis on ostcological grounds, the similarities it shows with
some of the beta anoles in chromosome structure are better in-
terpreted as a departure from the "standard" iguanid karyotype
that converged with some of the modifications shown in the anoles
of the grahami and chrysolepis series. Admittedly, the amount
of this convergence may be considerable. It would be possible to
derive the karyotype of A. jacare from that of A. chrysolepis by
centromeric fissions in the last pair of macrochromosomes of the
latter, leading to two pairs of microchromosomes with terminal
centromeres. This process would result in a complement with
six pairs of macrochromosomes separable into three distinct
groups, and in ten pairs of microchromosomes, exactly as in /I.
jacare. The osteological evidence, however, does not support any
close relationships between these two species.
The chromosome analysis thus indicates that Anolis jacare is an
alpha Anolis that has departed significantly from other members
of this group in chromosome number and structure, though main-
taining the standard iguanid karyotypic feature of six pairs of
macrochromosomes.
Observations in life (C. Rivero-Blanco and E. E. Williams):
Since no information of any ecological sort had ever been pro-
vided for Anolis jacare, it was as important an objective of the
expedition to Merida to provide this information as to obtain
chromosome data.
Only twelve anoles were collected during a period of three days
of active search. All were obtained on medium and large-sized
trees bordering the small Rio Milla just outside the city of Merida
(1639 meters above sea level). Several other areas within and
outside the city of Merida were carefully examined.
The general area is classified as Premontane Humid Forest in
the scheme of L. R. Holdridge (J. J. Ewel and A. Madriz. 1968).
The mean annual temperature is 19.1° C and the annual rainfall
1791 mm.
The two actual collection sites (Fig. 2) were roadside localities
and were subject to more or less penetration by the sun, especially
so in site 1 where trees were partly separated, less so in site 2
where the canopy was closed. In the first site, the anoles were
8
BREVIORA
No. 353
seen and collected on the branches of "majagua" (Heliocarpus
popoyensis, Tiliaceae) and "guamo" {Inga sp., Leguminosae), at
the second on "anime" {Montanea quadrangularis, Compositae)
and on a very large tree 10-15 meters high, not identified, since
leaves and flowers were not collected.
Figure 2. Map of the collecting sites for A nulls jacarc along the Rio
Milia outside Merida.
Collecting was done with the aid of a 5 meter long telescopic
fishing rod with a nylon noose. The animals were not shy but
avoided the noose by moving around the branch or further along
the branch or to other branches along the trunk or out on the finer
twigs. Two escaped high into the canopy; others did not move at
1970 ANOLIS JACARE 9
all. The number of animals seen varied from none on many trees
to four on one guamo tree. Few of those seen escaped the noose.
We have no belief that we have even the beginning of knowledge
of the population density of this species. The animals were difficult
to see and commonly lay along branches, and only twice were
they seen on the main trunk of the trees. They obviously ranged
widely within the trees they inhabited, including very high in the
crown. The first specimen taken came from a guamo tree that
was examined several times every day and even one night. It was
this tree that, on the last afternoon, provided three additional ani-
mals to give a final result of two males and two females on a
tree no more than six meters high and not especially complex.
This result was possible only because, during the last afternoon,
we had the help of a young local boy who was an excellent climber
and who was able to spot from a higher position animals that could
not be seen from below because of their resting position on
branches.
In summary, this is an animal inhabiting primarily the crown
and its branches, though not avoiding the trunk. It is not re-
stricted to shade; several individuals seen were in partial sun or
moved into sun without reluctance. It has no evident competitors.
No other lizards were seen in the collecting area either on the
trees or on the ground. Elsewhere in the vicinity other lizards
were found: Polychrus, in a hedge, and Aiueiva and Cnemido-
phorus, on the ground.
Discussion (E. E. WiUiams):
The karyotypic evidence clearly demonstrates a strong separa-
tion between jacare and either of the stocks of Lesser Antillean
anoles. Equally there is sharp difference between jacare and the
few mainland alphas that have been studied thus far (Gorman,
personal communication). On the face of the evidence, A. jacare
seems to occupy a rather isolated phyletic position.
It may be of interest and importance here that A . jacare is dis-
tributionally isolated also and that, very unusually for South
American anoles, it is not known to be sympatric with any con-
geners in any part of its known range.
There are other South American species that extend beyond
the range of their congeners somewhere at the periphery of their
range. A. jacare is special in that so far as known its whole range
is outside contact with any other anole.
Recent studies by T. Schoener (1970) have shown that in the
Lesser Antilles, "solitary" species, i.e., species without sympatric
10 BREVIORA No. 353
congeners, tend to be very similar in size and habitus. There also
appears to be a broadened unsp:ciaiized ecology characteristic
of these "solitary" anoles. We have noted above that the
bimaculatiis and roquet species groups are extraordinarily similar
in scale characters. We emphasize now that they are so in spite
of the fact that they are products of two quite separate invasions
of the West Indies and are very distinct in karyotype and bio-
ci'icm'stry.
Schoener infers, and we may agree with him, that some common
selective factor must be at work to keep (or evolve) external
similarity when wide underlying differences exist. That common
selective factor would appear to be the negative one of the absence,
or extreme limitation, of the number of congeners.
Certainly on the larger islands of the Greater Antilles a con-
trary rule exists: syntopic anoles are very diverse in morphology
or size or both.
The modification of a species in the absence of congeners or
other competitors in its general niche is sometimes spoken of as
"release." In morphology, at least, it is proper to speak of a more
positive selection than that implied by that essentially negative
term. A certain size seems clearly optimal and presumably the
features of squamation must likewise be held under selective
control.
In ecological behavior, "release" seems a more descriptive term,
since the wider range of habitat permitted a species in the absence
of close competitors concords better with our intuitive sense of the
meaning of release.
In the Lesser Antilles, there is often only one species per island
and, except for instances of very recent importation and their very
local occurrence (e.g., wattsi on St. Lucia, Underwood, 1959.
1962), there is a maximum of two species per island. These are
relatively old islands and the species on them are well differen-
tiated. They afford the classic and best examples of "solitary"
anoles.
A. jo.care, however, is as isolated in the Andes of Merida as the
solitary anoles of the Lesser Antillean islands. It is effectively on
a mainland island; it is interesting therefore, but not unexpected,
however, to find it resembling and behaving like an island anole —
a solitary anole of an old small island.
The resemblances, then, of A. jacare to A. leachii or A. luar-
moratus are to be explained in terms of adaptation to similar selec-
tive pressures. We need not, in fact, seek any complex zoogeo-
graphic solution to the similarity of one anole on island mountains
1970
ANOLIS JACARE
11
to one on a distant island; the similarity is non-phyletic, strictly
convergent.
TABLE
1
snout-vent
length of
adult -'
jacarc
73 mm
leach a
96 mm
intirnioraliis
77 mm
scales acioss
snout
6-8
4-5
4-5
scales between
semicircles
0-2
0-1
0-1
loreal rows
4-5
4-5
4-5
scales between
interparietal
and semi-
circles
1-3
1-2
1-2
supralabials to
center of eye
6-9
7-8
7-8
mental
number of
sublabials in
contact with
infralabials
not deeper
than wide
3-5
not deeper
than wide
2-4
deeper than
wide
2-4
scales between
sublabials in
contact with
mentals
4
4-6
3-4
ventrals
smooth
smooth
feebly keeled
lamellae under
phalanges ii and
iii of fourth toe
tail
dewlap
color
19-25
compressed
but without
crest, 2
dorsal rows
large
green with
variable dark
vermiculations
26-32
compressed,
with strong
crest in
males
small
24-30
Compressed, with
weak dorsal crest
in males
large
green with dark green with light
vermiculations vermiculations
but these on head in males
stronger on head only
than on body
12 BREVIORA No. 353
ACKNOWLEDGMENTS
The expedition to obtain and study A uoUs jaccire was supported
by the Instituto de Zoologia Tropical, Universidad Central de-
Venezuela, and by National Science Foundation Grant GB 6944
to Ernest E. Williams. Thanks are also due to Ingrid LcJbig for
help in laboratory work, and to Pedro Durand for facilities provided
at the Universidad de los Andes during the work in Mcrida. We
are indebted to George Gorman for communicating to us data on
the karyotypes of two female A . jacare.
REFERENCES
BouLENGER, G. A. 1903. On some batrachians and reptiles from
Venezuela. Ann. Mag. Nat. Hist., ser. 7. 11: 481-484.
Etheridge, R. 1960. The relationships of the anoles ( Reptilia:Saiiria:
Iguanidae) an interpretation based on skeletal morphology. Ann Arbor,
Michigan: University Microfilms, xiii -f 236 pp.
Ewel, J. J., AND A. Madriz. 1968. Zonas de Vida de Venezuela.
Ediciones del Fondo Nacional de Investigaciones Agropecuarias.
Caracas. 265 pp + Mapa Ecologico de Venezula.
Gorman, G. C. 1965. Interspecific karyotypic variation as a systematic
character in the genus Anolis (Sauria: Iguanidae). Nature 208: 95-97.
Gorman, G. C, and L. Atkins. 1966. Chromosomal heteromorphism
in some male lizards of the genus Anolis. Amer. Nat. 100: 579-583.
1967. The relationships of the Anolis of the roquet species
group (Sauria; Iguanidae). II. Comparative chromosome cytology.
Syst. Zool. 16: 137-143.
1968a. New karyotypic data for 16 species of Anolis
(Sauria: Iguanidae) from Cuba. Jamaica and the Cayman Islands.
Herpetologica 24: 13-21.
1968b. Confirmation of an X-Y sex determining mech-
anism in lizards (Anolis). Copeia 1968: 159-160.
1968c. Natural hybridization between two sibling species
of Anolis lizards: chromosome cytology. Science 159: 1358-1360.
1969. The zoogeography of Lesser Antillean Anolis
lizards — an analysis based upon chromosomes and lactic dehydro-
genases. Bull. Mus. Comp. Zool. 138: 53-80.
Gorman, G. C, L. Atkins, and T. Holzinger. 1967. New karyotypic
data on 15 genera of lizards in the family Iguanidae, with a discus-
sion of taxonomic and cytological implications. Cytogenetics 6:
286-299.
1970 ANOLIS JACARE 13
Gorman. G. C. R. Thomas, and L. Atkins. 1968. Intra- and inter-
specific chromosome variation in the lizard Anolis cristate I Ins and its
closest relatives. Breviora 293: 1-13.
Levan, a.. K. Fredga. and A. A. Sandberg. 1964. Nomenclature for
centromeric position on chromosomes. Hereditas 52: 201-220.
Schmidt, K. P. 1939. A new lizard from Mexico with a note on the
genus Norops. Zool. Ser.. Field Mus. Nat. Hist. 24: 7-10.
ScHOENER, T. W. 1970. Size patterns in West Indian Anolis Lizards. II.
Correlations with sizes of particular sympatric species displacement
and convergence. Amer. Nat. 104: 155-173.
Underwood, G. 1959. The anoles of the Eastern Caribbean (Sauria,
Iguanidae). Part III. Revisionary notes. Bull. Mus. Comp. Zool.
121: 187-226.
___. 1962. Reptiles of the Eastern Caribbean. Caribbean
Affairs (N.S.) 1: 1-192.
(Received 17 April 1970.)
14 BREVIORA No. 353
A^ A2 A3 ^ -*\ » U ^
XX XX .. ^^W
B1 B2 CI
• • •
Figure 3. Spermatogonia! metaphase and karyotype of AiioUs jacarc.
Specimen no. MCNC 5601. cell no. A-167 T5 C2. Scale: 10 micra.
M }t n «V^
• •
« »
Figure 4. Spermatogonial metaphase and karyotype of Aiiolis jacare.
Specimen no. MCNC 5604. cell no. A-)71 T.^ C2. Scale: 10 micra.
970
ANOLIS JACARE
LS
w*-
Figure 5. Diakinesis of Aiuilis hicarc. Specimen no. MCNC 5603, eel
no. A-170 T7 CI.
• ..^ \^IX-''
^■^-rj.^*^
W~'f'^'
-^ml
^:
■%,.
~ "^'
j^;;.-.k.
Figure 6. Rio Milla. Merida (site 1 of Fig. 2). To the right of the hght
pest is the guamo (Inga sp.) in which two males and two females were
collected. The other trees to the right are majagua (Heliocarpus
pupayensis) where other specimens were collected
BREVIORA
Musenaiiii of Compsirative Zoology
Cambridge, Mass. 18 September, 1970 Number 354
TAXONOMIC AND ECOLOGICAL NOTES ON SOME
MIDDLE AND SOUTH AMERICAN LIZARDS OF THE
GENUS Ameiva (TEIIDAE)
Arthur C. Echternacht
Abstract. The taxonomy of two Middle American and one South
American Ameiva (Sauria, Teiidae) is discussed. Ameiva festiva niceforoi
Dunn is accorded species rank, A. f. miadis Barbour and Loveridge is for-
mally designated a subspecies of A. undiilata. and A. iiiuliilata thomasi
Smith and Laufe is placed in the synonymy of A. chaitzami Stuart. Diag-
noses and statements of range are provided for each, and ecological in-
formation is presented for miadis. The condition of the median parietal
(divided or not) is shown to be unstable in Ameiva and useless for diag-
nosing species in Middle America.
INTRODUCTION
In the course of my studies of geographic variation in the
Middle American species of the lizard genus Ameiva, I have found
that certain taxa require reallocation and that diagnoses presented
with the original descriptions of some are either in error or mis-
leading. It is the purpose of this paper to clarify the taxonomic
positions of three of these taxa. A diagnosis is presented for
each, and ecological information is included where warranted.
Acknowledgements. I am indebted to the following persons
for the loan of specimens in their care: W. E. Duellman, Univer-
sity of Kansas Museum of Natural History (KU); R. F. Inger
and H. Marx, Field Museum of Natural History (FMNH); E. V.
Malnate, Academy of Natural Sciences of Philadelphia (ANSP);
J. R. Meyer (Private Collection); J. A. Peters and G. R. Zug,
United States National Museum (USNM); C. F. Walker and L.
C. Stuart, University of Michigan Museum of Zoology (UMMZ);
R. G. Zweifel, American Museum of Natural History (AMNH);
2 BREVIORA No. 354
and E. E. Williams, Museum of Comparative Zoology (MCZ).
W. E. Duellman and E. E. Williams read and criticized the manu-
script.
Ameiva niceforoi Dunn
Ameiva f estiva niceforoi Dunn, Notulae Naturae no. 126: 1-2, 1943
(Holotype: ANSP 24300. Type Locality: "Sasaima, in the eastern Andes,
75 km northwest of Bogota, altitude 1200 meters," Colombia. Collector:
Hermano Niceforo Maria).
Diagnosis. Ameiva niceforoi can be distinguished from its
congeners by the following combination of characters: Small size
(maximum observed snout-vent length |SVL] 82 mm for males,
75 mm for females) ; central gular scales much enlarged, surround-
ing scales diminishing in size gradually toward the periphery of
the gular region; frontal and frontoparietal scales entire; three
parietal scales; total number of femoral pores moderate; number
of dorsal granules around the body (GAB) and occiput to rump
(GOR) low; no preanal spurs; broad middorsal stripe bordered
laterally by a fine white stripe; black dorsolateral stripe bordered
ventrally by a fine white stripe; no narrow, light-colored vertebral
stripe.
Range. Knovv'n only from the type locality and from Honda,
Departamento Tolima, Colombia.
Remarks. Dunn (1943) diagnosed Ameiva f estiva niceforoi
( =A. niceforoi) as "A form of f estiva, identical with it in size,
proportions, and in scalation, but remarkably different in mark-
ings." In addition to color pattern (Fig. lA), niceforoi differs
from fesiiva in a number of characters of scutellation. In color
pattern, niceforoi is virtually identical to female or subadult male
A. leptophrys but differs from leptoplvys in numerous scale char-
acters. Some pertinent differences among the three species are
summarized in Table 1. Because geographic variation is marked
(Echternacht, 1970), means of characters of scutellation for an
entire species tend to mask similarities between niceforoi and
samples of other species drawn from nearby localities. For this
reason, means for nearby samples are given as well as those for
the entire species. In addition to the total number of femoral
pores, GAB and GOR, niceforoi differs from leptophrys in the
arrangement of scales peripheral to the enlarged central gulars. In
leptoplvys the posterior gular scales are much reduced in size
1970
NOTES ON AMEIVA
Figure 1. (A) Ameiva niceforoi, paratype (ANSP 24303: Sasaima,
Depto. Cundinamarca, Colombia), snout-vent length 77 mm. (B) Ameiva
undulata miadis, holotype (MCZ 26970: Isla del Maiz Grande, Depto.
Zelaya, Nicaragua), 126 mm. (C) Ameiva chaitzami, paratype (MCZ
52170 [formerly UMMZ 90642]: Along Cahabon-Languin trail ca. 2 km
N Finca Canihor, Depto. Alta Verapaz, Guatemala), 69 mm. All males.
relative to the anterior gulars. Ameiva niceforoi and festiva are
similar in this respect.
No other South American species of Ameiva seems to be closely
related to any of the three species discussed above. An Ameiva
similar to niceforoi could have given rise to either leptophrys or
festiva or to both, but the present chaotic situation with respect
to the taxonomy of Ameiva in South America (Medem, 1969) pre-
cludes decisive conclusions concerning phylogenetic relationships.
Studies in progress are designed to clarify this situation.
Other than the type series (ANSP 22784, 24300-303), only
three specimens of Ameiva niceforoi are known: USNM 93500-
93501 (Topotypes) and AMNH 35300 from Honda, Depto.
Tolima, Colombia.
BREVIORA
No. 354
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1970 NOTES ON AMEIVA 5
Ameiva undulata miadis Barbour and Loveridge
Ameiva f estiva miadis Barbour and Loveridge, Bull. Mus. Comp. Zool.,
69: 141-142, 1929 (Holotype: MCZ 26970. Type Locality: "Great Corn
Island" [=Isla del Maiz Grande], Depto. Zelaya. Nicaragua. Collector:
James L. Peters).
Ameiva undulata miadis, Dunn, Proc. Acad. Nat. Sci. Philadelphia, 92:
115, 1940. Villa, Rev. Biol. Trop., 15: 119, 1968.
Diagnosis. Ameiva undulata miadis (Fig. IB) can be dis-
tinguished from A. f estiva and other subspecies of A. undulata
on the basis of its unique color pattern: No narrow, light-colored
vertebral stripe; lateral pattern of blue (males) or gray-brown
(females) bars on a black background extending from mid-
dorsal stripe to enlarged ventral scutes; broad middorsal stripe
from occiput onto tail, much disrupted by transverse black blotches
that are often continuous with the black background color of the
flanks; sexual dimorphism in color, but not pattern, in adults.
Range. Known only from Islas del Maiz, Depto. Zelaya, Nica-
ragua.
Remarks. With the exceptions noted above, Ameiva undulata
miadis is similar to other subspecies of undulata that have barred
flanks (see Echternacht, 1970). Dunn (1940) and Villa (1968)
have briefly alluded to the relationship of miadis with undulata
Barbour and Loveridge (1929) considered Ameiva festiva
miadis ( ^A. undulata miadis) to be intermediate between A.
festiva and A. ruthveni ( =A. leptophrys) . Color pattern alone
distinguishes miadis from leptophrys, but in addition, the peri-
pheral gular scales of undulata are like those described for niceforoi
above and differ from those of leptophrys in the same manner.
Also, in leptophrys the postnasals are not in contact with the pre-
frontal scales, whereas in undulata these scales are in contact.
The nearest known mainland populations of Ameiva festiva are
at Bluefields, Depto. Zelaya, Nicaragua, about 85 kilometers west
of Isla del Maiz Grande. Ameiva festiva from Bluefields are
typical of the species in having a well-defined vertebral stripe in
ah but large adults and in having little barring on the flanks.
Nearest known populations of leptophrys are in the southeastern
lowlands of Costa Rica, and the nearest populations of undulata
are in northeastern Honduras, western Nicaragua, and northwest-
ern Costa Rica. Four specimens of undulata (ANSP 15438-
15441) from Huaunta Haulover, Depto. Zelaya, Nicaragua, about
BREVIORA
No. 354
150 kilometers north-northwest of Isla del Maiz Grande, may in-
dicate an isolated population there. In color pattern, the undulata
from Islas del Maiz most closely resemble those from near Cata-
camas, Depto. Olancho, Honduras. In these Honduranian
Ameiva, the middorsal stripe is heavily blotched, but the blotches
are never continuous onto the flanks.
Island populations of Middle American Ameiva (with the ex-
ception of qiiadnlineata) tend to have a relatively high GAB count
when compared to mainland samples of the same species (see
Echternacht, 1970). Although the mean for this character does
not differ statistically from those of certain mainland samples
examined, it is the highest recorded for the species {A. undulata:
X = 149.5, standard deviation of the mean = 16.5, range =
108-202, N = 918. A. u. miadis: x = 186.3 ± 9.3, range =
165-202, N = 42).
Nothing has been reported of the ecology of Ameiva undulata
miadis. I was able to make some observations on June 4-8, 1966.
The lizard is very common on Isla del Maiz Grande and I found it
particularly abundant in clearings within the coconut groves that
cover the island (Fig. 2). It was also common in the shrub and
'i.. -^ *t,. ., ,.^5^-s.
Figure 2. Habitat of Ameiva undulata miadis on Isla del Maiz Grande,
Depto. Zelaya, Nicaragua. June 1966.
1970 NOTES ON AMEIVA 7
grassy vegetation along the airstrip and in clearings around build-
ings. None were seen on open beaches. Individuals often were
encountered basking or foraging on or around fallen palm fronds.
Activity extended from dawn until dusk, but more lizards were
seen early in the day than in late afternoon. The lizards were not
active in rain. Hatchlings with clearly dehneated umbilical scars
were observed. The chin, gular region, and anteriormost part of
the chest of many males was bright reddish orange. In others,
this area was pale blue. Such dichromatism is widespread among
species of Ameiva and may be associated with reproductive con-
dition. Evidence is lacking on this point. English-speaking resi-
dents of the island refer to the Ameiva simply as "lizard," al-
though they have specific names for Iguana iguana and Cteno-
saura similis, both of which are common.
Ameiva chaitzami Stuart
Ameiva chaitzami Stuart, Proc. Biol. Soc. Washington, 55: 143, 1942
(Holotype: UMMZ 90638. Type Locality: "Along Cahabon-Languin trail
about 2 km north of Finca Canihor . . . , Alta Verapaz, Guatemala." Col-
lector: L. C. Stuart).
Ameiva imdulata thomasi Smith and Laufe, Univ. Kansas Sci. Bull., 31:
47-50, pi. lA, 1946 (Holotype: FMNH 100006. Type Locality: "La Liber-
tad, Chiapas, near Rio Cuilco where it crosses the Guatemalan border,"
Mexico. Collector: Henry D. Thomas). New synonymy.
Diagnosis. Ameiva chaitzami (Fig. IC) can be distinguished
from its congeners by the following combination of characters:
Small size (maximum observed SVL 85 mm for males, 75 mm for
females); central gular scales enlarged, in longitudinal series;
gradual reduction in size of scales radiating outward from the
central gulars; prefrontals in contact with postnasals; three parietal
scales (four if median parietal divided); transverse row of abrupt-
ly enlarged mesoptychial scales; usually eight transverse rows of
ventral scutes at midbody; moderately narrow middorsal stripe
(mean width in terms of granules 37.1, N = 61); no narrow,
well-defined vertebral stripe; often a dark secondary stripe medial
to the dorsolateral light stripes; dorsolateral blotches of adult
males fused to the dorsolateral light stripe so that the latter has
a well-defined dorsal border, an irregular ventral border. The
latter character alone will distinguish chaitzami from undulata
with certainty.
8 BREVIORA No. 354
Range. Valleys of the upper tributaries of the Rio Grijalva
in Chiapas, Mexico, and west-central Guatemala, from the vicin-
ity of Finca Canihor, Depto. Alta Verapaz, Guatemala, and near
Poptiin, Depto. El Peten, Guatemala.
Remarks. Stuart (1942) diagnosed Ameiva chaitzami as "An
Ameiva almost identical with Ameiva nndulata stuarti Smith from
which it may readily be distinguished by the fact that the median
parietal is divided longitudinally to produce four instead of three
parietals." The posterior scales on the dorsal surface of the head
is an unstable character among Middle American species of
Ameiva, and diagnoses based on scales in the area are unreliable
(Echternacht, 1970). Considering nndulata only, I noted the
median parietal to be divided or semidivided in 22.4 per cent of
1043 specimens obtained from throughout the range of the species.
This is probably a conservative estimate of the frequency of oc-
currence of this condition, because the information was not noted
early in the study. There seem to be no geographic trends as-
sociated with the condition of the median parietal. At least 38
of 45 samples contained individuals with divided or semidivided
median parietals. Samples having a high frequency of occurrence
of division were as follows: near Chinandega, Depto. Chinandega,
Nicaragua (55 per cent, N = 31); Piedras Negras, Depto. El
Peten, Guatemala (72 per cent, N = 32); Canihor, Depto. Alta
Verapaz, Guatemala (50 per cent, N = 30); near Panajachel,
Depto. Sololii, Guatemala (48 per cent, N = 29); Sabana de
San Quintin, Chiapas, Mexico (100 per cent, N = 10); near Las
Tazas and Florida, Chiapas, Mexico (90 per cent, N = 30).
Three samples of chaitzami yielded the following frequencies:
Comitan, Chiapas, Mexico, 3 per cent (N = 30); near San An-
tonio Huista, Depto. Huehuetenango, Guatemala, 10 per cent
(N = 31 ); near Finca Canihor, Depto. Alta Verapaz, Guatemala,
and near Poptun, Depto. El Peten, Guatemala, 89 per cent (N =
9). Six of the nine in the last sample constitute the type series.
At the type locality and at Poptun, chaitzami is sympatric with
A . imdidata hartwegi Smith, a large subspecies quite distinct in color
pattern and scutellation from chaitzami. The samples from Canihor
and Piedras Negras, Guatemala, and from Sabana de San Quintin,
Chiapas, are hartwegi. Elsewhere within its range, samples of
hartwegi have from 13 to 30 per cent of individuals with divided
or semidivided median parietals. It is noteworthy that the holo-
type of hartwegi (FMNH 108600), obtained across the Rio Usu-
macinta from Piedras Negras, has a divided median parietal.
1970 NOTES ON AMEIVA 9
Smith and Laufe (1946) discussed the evolution of Aineiva
undidata and recognized several new subspecies, but they ap-
parently did not examine specimens of chaitzami. The description
of A. u. thomasi (Smith and Laufe, 1946) agrees with that of
Stuart (1942) for chaitzami in most respects. I have examined
the type specimens of both and consider them to be conspecific.
The samples cited above from Comitan and near San Antonio
Huista are from within the range of thomasi as described by
Smith and Laufe (1946).
LITERATURE CITED
Dunn, E. R. 1940. New and noteworthy herpetological material from
Panama. Proc. Acad. Nat. Sci. Philadelphia, 92: 105-122.
. 1943. A new race of Ameiva f estiva from Colombia. Notu-
lae Naturae no. 126, 2 pp.
EcHTERNACHT, A. C. 1970. A review of Middle American lizards of the
genus Ameiva (Teiidae) with emphasis on geographic variation. Un-
published Ph.D. Dissertation, University of Kansas, Lawrence, Kansas.
Medem, F. 1969 (1968). El desarrollo de la Herpetologia en Colombia.
Rev. Acad. Colombiana Cienc. Exactas Fis. Natur., 13(50): 149-199.
Barbour, T., and A. Loveridge. 1929. Vertebrates from the Corn Is-
lands. Reptiles and Amphibians. Bull. Mus. Comp. Zool., 69: 138-146.
Stuart, L. C. 1942. Comments on the iindiilata group of Ameiva
(Sauria). Proc. Biol. Soc. Washington. 55: 143-150.
Smith, H. M., and L. E. Laufe. 1946. A summary of Mexican lizards
of the genus Ameiva. Univ. Kansas Sci. Bull., 31(2): 7-73.
Villa, J. D. 1968. A new colubrid snake from the Corn Islands, Nic-
aragua. Rev. Biol. Trop., 15: 117-121.
(Received 13 June 1970.)
BREVIORA
Miasemim of Coimpsirsitive Zoology
Cambridge, Mass. 30 November, 1970 Number 355
GENERIC RELATIONS AND
SPECIATION PATTERNS IN THE CARACARAS
(AVES: FALCONIDAE)
Francois Vuilleumier^
Abstract. The caracaras are a group of American Falconidae occurring
from temperate and subtropical North America southward to extreme south-
ern South America and the Falkland Islands. The taxa of caracaras appear
to be closely interrelated. It is suggested that they be classified in two
genera: Daptrius (forest caracaras; two sympatric species), and Polybonis
(nonforest caracaras; two species-groups: the chimachima and planciis
species-groups, with two and three species, respectively). Former classifica-
tion advocated the use of four genera. Speciation is long completed in
Daptrius and in the Polybonis chimachima species-group. In the Polybonis
planciis species-group, however, several phenomena are evidence of active
species formation. Geographical isolates exist that are morphologically
differentiated enough to be considered borderline cases between species and
subspecies. Some of the cases of geographical isolation in the caracaras can
be related to climatic and vegetational changes following glacial events of
the Pleistocene.
INTRODUCTION
This paper constitutes the fourth of a series stemming from
studies on speciation in Andean birds (see Vuilleumier, 1968,
1969, 1970).
I shall discuss, first, the generic classification of the caracaras,
and secondly, the patterns of distribution, geographical variation
and speciation in these birds, with particular emphasis on the
Andean taxa. I shall deal either with problems not, or only litde,
covered by Brown and Amadon (1968) in their recent book, or
1 Biology Department, University of Massachusetts, 100 Arlington Street,
Boston, Massachusetts 02116.
2 BREVIORA No. 355
with controversial issues, especially when my own conclusions differ
from theirs.
The data were obtained from examination of about 250 skins
and some skeletons; and from field studies on the habitat prefer-
ences, general behavior, and distribution of five taxa of caracaras
made over a period totaling thirteen months during trips to South
America in 1964, 1965, and 1967-68.
THE CARACARAS
The 7 to 10 or 11 species of caracaras can be distinguished
from other Falconidae more by their vulture-like external morpho-
logical characters and associated scavenging habits, than by clear-
cut anatomical characters (Friedmann, 1950: 719). They differ
from true falcons, however, in their habit of building their own
nest, as pointed out by Brown and Amadon (1968: 23, 104). The
problem of whether the caracaras should be accorded taxonomic
rank within the Falconidae, and if so, which one (e.g., subfamily,
tribe), will not be discussed here.
The species of caracaras have traditionally been placed in four
genera (see, e.g., Peters, 1931; Hellmayr and Conover, 1949;
Friedmann, 1950; de Schauensee, 1966; Brown and Amadon,
1968): Daptrius Vieillot, 1816, with two species; Milvago Spix,
1824, with two species; Phalcoboenus d'Orbigny, 1834, with two
to four species; and Poly bonis Vieillot, 1816, with one to three
species. (The long controversy over the names Polyborus versus
Caracara Merrem, 1826, was resolved by Amadon (1954), who
showed that "Vieillot's diagnosis of Polyborus applies to the Cara-
caras," so that "the diagnosis sustains the name." Caracara must,
therefore, go into synonymy, and the issue can be considered
closed.)
As a group, the caracaras are distributed from temperate North
America southward to Central and South America, ranging as far
south as the Falklands and the islands off Tierra del Fuego. Sev-
eral species have very broad ranges, especially Polyborus plancus,
the distribution of which encompasses almost that of the entire
group. Other species are, on the contrary, quite localized geo-
graphically. For example, the extinct Polyborus lutosus occurred
only on Guadalupe Island off Baja California, and the living Phal-
coboenus australis breeds on a handful of small islands off the
southern coast of Tierra del Fuego and on the Falklands.
1970 SPECIATION IN THE CARACARAS 3
Correlated with the broad distribution of the caracaras is their
ecological diversity. As a group, these birds inhabit most vegeta-
tion formations found in their geographical range, from desert
scrub to tropical lowland wet forest, and including the highest
zones of Andean vegetation. About the only major type of vegeta-
tion not favored by caracaras is montane tropical wet forest (or
cloud forest).
All the taxa of caracaras are scavengers, but should perhaps be
viewed as omnivorous, since their diet also includes live prey (ver-
tebrates and invertebrates) and vegetal matter (see, e.g., Hud-
son, 1920: 62-88; Wetmore, 1926: 92-96; Haverschmidt, 1962:
157-158; Friedmann, 1927: 157; Friedmann and Smith, 1950:
450-451, and 1955: 486-487; Brown and Amadon, 1968). Among
the most notable food specializations of the caracaras is the habit
of Polyborus planciis of attacking domestic animals the size of
sheep (Johnson, 1965: 263), and the marked predilection shown
by Daptrius americanus for colonial wasps (Skutch, 1959).
Most caracaras are social, at least during the nonbreeding sea-
son, and form intraspecific flocks that appear to be feeding associa-
tions. The larger-sized species may form smaller flocks than the
smaller ones. Thus the largest flock of the osprey-sized Polyborus
plancus I have observed comprised about 15 birds, while I have
often seen much larger flocks of the kestrel-sized Milvago chimango.
The intraspecific gregariousness of some of the smaller species may
also extend to the breeding season. Drury (personal communica-
tion ) observed a colony of Milvago chimango where the nests were
only about 30 feet (ten meters) apart. The caracaras also form
associations with vertebrates other than birds. In Patagonia, Mil-
vago chimango flocks are frequent near cattle and horses; I even
saw one bird sitting on the flank of a lying horse, pecking from time
to time at the skin, perhaps to eat ticks. Hudson (1920: 70) men-
tioned that M. chimango "follows the plough," thus playing the
part of gulls {Lams) elsewhere. The habit of following moving
vehicles from which scraps of food can be collected has been
observed in Phalcoboenus megalopterus along the Yungas Road
in La Paz, Bolivia (Niethammer, 1953: 265; personal observa-
tion), and in Polyborus plancus along a railroad in the Bolivian
Chaco (Eisentraut, 1935: 391). The gregarious behavior exhib-
ited by the caracaras may be correlated to a large extent with their
scavenging habits, yet, from my own observations, I would judge
their behavior to be much more plastic and diverse than that of
4 BREVIORA No. 355
other scavengers such as Old World or New World vultures
(Cathartidae and Aegypiinae, respectively).
ANALYSIS OF SOME CHARACTERS OF THE CARACARAS
Size
The caracaras vary considerably in size, as several authors
(Friedmann, 1950; Brown and Amadon, 1968) have already
pointed out. This variation is obvious if wing length, in the ab-
sence of data on weights, is used as an indicator of overall body
size (Table 1). Taxonomists have often been bothered by size
differences between species, and have been reluctant to place in
the same genus closely related species which differed conspicuously
in size but in few, or no, other characters. This attitude might
have been prevalent among the ornithologists who worked with
caracaras, because the two smallest species belong in the genus
Milvago and the largest species in Phalcoboenus and Polyborus.
Yet the two species of Daptrius bridge the gap between these ex-
tremes. I therefore believe that size should not be given undue
weight in the supraspecific classification of the caracaras. In other
Falconidae, notably in the genus Falco, similarly large size differ-
ences between species have not prevented their inclusion in the
same genus.
Proportions
In a number of bird taxa, a proportionately short tarsus is cor-
related with arboreal habits, and a long tarsus with more terrestrial
habits. Most species of caracaras are both arboreal and terrestrial,
but some definite trends toward one or the other of these habits
exist, especially in regard to feeding habits. Thus Polyborus and
Milvago are often seen perched on trees, and they breed in trees,
but they do a lot, perhaps most, of their foraging for food on the
ground. The species of Phalcoboenus (with the possible exception
of P. albogularis) seem to forage entirely on the ground, and breed
in cliffs and rocky slopes; they do on occasion perch on buildings.
The species of Daptrius seem at variance with the other three
genera because they do some, perhaps even a substantial, portion
of their foraging for food in trees.
From this summary one would thus expect the species of Dap-
trius, which are more arboreal than the other caracaras, to have
proportionately shorter tarsi than other species. Figure 1 shows a
1970
SPFXIATION IN THE CARACARAS
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60
6 BREVIORA No. 355
scatter diagram of tarsus length plotted against wing length (rep-
resenting body size in the absence of weight data) in the caracaras.
The two species of Milvago, although overlapping considerably in
wing length, show little overlap in tarsus length. The southern
species, M. chimango, is decidedly longer-legged than the northern
one, M. chimachima. This difference, especially in regard to the
zone of sympatry of the two species, is discussed again below in
the section on speciation in these birds. Dapthiis ater has somewhat
longer wings than either species of Milvago, but its tarsal measure-
ments overlap fully with those of M. chimachima. Yet D. ater
would seem more arboreal than M. chimachima. The arboreal
Daptrius americanus overlaps with the more terrestrial Phalco-
boenus megaloptenis, P. albogularis, and Polyborus plancus in
wing length, but has a much smaller tarsus. In this case the differ-
ence in tarsus size appears to correspond to a difference in habits.
TARSUS LENGTH Cmm)
\
-•■□ • Y-^-:
I* * ,'
\*_.. -■■-'■
*;
/ 4 * «"*> /
.Q' i®.
WINO LENSTM (mm)
(*j^'' planeus
:*;:,
\*^*^ chimango /odj megalopi er u«
5JP.S-'
\P^/' ater '\**y "Ibogulari*
\^i-. amerieanuo
Figure 1. Tarsus length plotted against wing length in seven taxa of
caracaras.
1970 SPECIATION IN THE CARACARAS 7
It is worthy of note that the species of Phalcoboemis closest to D.
americamis in tarsus length is P. albogularis (see Fig. 1) said by
Olrog (1948: 478; 1950: 520) to occur in forests and thus to
diverge from the habitat preferred by P. megalopterus.
On the basis of the proportions of tarsus and wing lengths plotted
on Figure 1 , Milvago chimango, M. chimachima, and Daptrius ater
would form one group of species closely similar to each other, over-
lapping greatly in both wing and tarsus length. A second such
group would include Phalcoboemis and Polyborus, while Daptrius
americanus appears clearly distinct from all other species, yet inter-
mediate between the two groups.
Interspecific variation in tail length is summarized in Table 2.
When handling skins, Daptrius americamis appears distinctly longer
tailed than the other species of caracaras. If the tail/wing ratios,
instead of the absolute tail lengths, are compared among the species
of caracaras, then a smooth variation is detected, from the propor-
tionately short-tailed Polyborus plancus and Phalcoboemis mega-
lopterus to the long-tailed Milvago chimachima and Daptrius
americamis. No groups of caracaras can be established on the
basis of relative tail length.
TABLE 2
Variation in tail length, expressed in percent of wing length,
among the caracaras. The taxa are ranked in order of increasing
tail/wing ratio. Both sexes, and adult and subadult birds included.
Taxon
Tail/ Wing
X 100
Sample Size
Range
Mean
plancus
50.8-58.1
53.9
42
megalopterus
53.1-57.4
55.5
13
carunculatus
55.6
1
chimango
54.8-58.7
57.3
5
albogularis
54.1-64.7
58.4
5
australis
57.5-61.5
59.4
4
ater
58.4-63.5
61.1
6
lutosus
62.7
1
chimachima
60.9-70.2
66.2
25
americanus
62.2-74.6
68.4
36
8 BREVIORA No. 355
Plumage Color and Pattern
Good descriptions of both adult and immature plumages of
caracaras have been given by Friedmann (1950) and Brown and
Amadon (1968), who should be consulted for full details. The
main colors of aduh and immature plumages are summarized in
Table 1.
Adult caracaras vary from a brown, rather unpatterned plumage
(such as is found in a variety of birds of prey) to a highly pat-
terned plumage. Thus Milvago chimango has a brown plumage,
variegated somewhat with barrings, reminiscent of the plumage of
some kites of the genus Milvus. The plumages of both species of
Daptrius, and of the three Andean species of Phalcoboenus (carun-
culatus, megalopterus, and albogularis) are very different from
those of the other caracaras. These birds are strikingly marked
with black and white patterns, and remind one of gallinaceous
birds such as Crax.
Immature caracaras are less varied in their plumages. Most of
them are brownish with patterns of barring or streaking, with the
exception of the immature Daptrius americanus, which is like the
adult. The dimorphism between immature and adult can be very
well marked, as in Phalcoboenus carunculatus, P. megalopterus,
and P. albogularis; or poorly marked, as in Daptrius americanus
or Milvago chimango; or intermediate, as in Polyborus plancus and
P. lutosus.
In spite of the variation in plumage color and pattern exhibited
by the different species of caracaras, certain basic resemblances are
evident in the group as a whole when adult and immature plumages
are compared together. For example, the immatures of Milvago
chimachima, Phalcoboenus carunculatus, P. megalopterus, P. albo-
gularis, Polyborus lutosus and P. plancus are all very similar to
one another, as they are to the adult of Milvago chimango. Thus,
regardless of the intraspecific variabihty between immatures and
adults on the one hand, and the interspecific variability among
adults on the other, Milvago, Phalcoboenus, and Polyborus all
appear to have similarities in plumage color and pattern.
In every genus except Polyborus, there is a dichotomy between
one or more species that have different immature and adult plum-
ages, and one or more species that have quite similar immature and
adult plumages. This dichotomy is obvious in Milvago, with
1970 SPECIATION IN THE CARACARAS 9
chimanfio plumages much alike, and chimachima unlike; in Phal-
coboenus, with australis plumages much alike, and with canincu-
latus-megalopterus-albogularis plumages very different. The dicho-
tomy is still present, but much less obvious, in Daptrius, with
americanus plumages alike and ater plumages slightly different.
Correlation between this dichotomy and patterns of geographical
distribution among closely related species is not readily apparent.
In Milvago and Phalcoboemis , the species which differ least in their
adult and immature plumages are the two southern ones; but in
Milvago alone, there is an important geographical overlap between
chimango and chimachima, whereas there is no or only a very nar-
row overlap between the southern australis and the more northern
carunculatus-megalopterus-albogularis complex. In Daptrius (di-
chotomy poorly marked), there is almost complete overlap between
the ranges of the two species, since the range of ater is almost
entirely contained within that of americanus. In Polyborus (no,
or almost no dichotomy ) , lutosus and plancus are entirely allopat-
ric. (The possible ecological significance of the immature and adult
dimorphism is discussed below under habitat preferences.)
If one were to consider the immature-hke plumage of some sex-
ually mature birds as being a primitive condition, and the strikingly
different plumage in other mature birds as an advanced one (i.e.,
secondarily developed during the evolutionary history of the cara-
caras), then Polyborus and Daptrius might be thought to be more
primitive than the other two genera. If a brownish, barred and
streaked immature plumage were considered a more primitive con-
dition, then Daptrius is more advanced in this character than the
three other genera, because in that genus, immatures are hardly
different from adults, being also conspicuously patterned in black
and white.
Naked Facial Skin
Every species in the genera Polyborus, Phalcoboenus, and Dap-
trius has brightly colored naked skin between the bill and the eye,
and often around the eye or even the throat. In the genus Milvago,
only M. chimachima has naked facial skin; M. chimango has a fully
feathered head. The area of unfeathered skin varies from species
to species among those that have naked facial skin. In the species
of Phalcoboenus, this variation is geographical. In Phalcoboenus
carunculatus (Colombia and Ecuador), the surface of facial skin
is extensive, and the throat is even adorned by fleshy wattles. In
10 BREVIORA No. 355
p. megalopterus (Peru, Bolivia, and northern Chile-Argentina),
only the lores are unfeathered, while the throat is largely feathered.
In P. albogularis (Patagonia), the extent of bare loral skin is
smaller than in megalopterus. Finally, in P. australis (southernmost
islands of South America), the facial skin is almost entirely feath-
ered, but there are wattles on the breast instead. The difference
between P. australis and P. carunculatus-megalopterus-albogularis
may be related to the fact that australis is sympatric with Polyborus
plancus, a species having extensive naked facial skin, whereas the
other three species of Phalcoboenus are the only caracaras in their
respective ranges.
The naked facial skin in the caracaras varies from salmon pink
to rose-red and from yellowish to bright red (see Table 1). This
variation is both intra- and interspecific. I have seen the facial
skin of one individual of Polyborus plancus change, in a few sec-
onds, from pale yellow to salmon-pink and finally to vivid red.
Such a rapid change cannot easily be ascribed to hormonal influ-
ences, but is more likely to be due to a sudden flush of blood to the
superficial vessels of the skin. Brown and Amadon (1968: 738)
mention a reverse change in P. plancus: "bare facial skin carmine
red, changing to yellow when excited."
The variation in facial skin color in the caracaras (Table 1)
seems too extensive within species, and too restricted between
species (differences between, say, yellow and orange, or orange and
red seem relatively slight) to be important as a species-specific
means of recognition. Presence or absence, together with color
and area of unfeathered skin might, however, play such a role
among true sympatric species, e.g., Phalcoboenus australis and
Polyborus plancus already cited, or Milvago chimango and M.
chimachima.
Nostril Shape
I have examined skulls of Polyborus plancus (several speci-
mens), Phalcoboenus australis (4), Daptrius sp. (1 ), and Milvago
sp. (3). Polyborus plancus has slanted, elongated nostrils (bean-
shaped), while Phalcoboenus australis, Daptrius sp., and Milvago
sp. all have rounded nostrils. This difference is also visible on the
cere of study skins and seems to be confined to Polyborus (Table
1 ) , since only P. lutosus and P. plancus have bean-shaped nostrils.
Swann (1925: 66) and Friedmann (1950: 545-546) used this
1970 SPECIATION IN THE CARACARAS 11
difference in nostril shape, together with other characters, as diag-
nostic features to key out the genera of caracaras. I do not know
what the biological significance of this difference may be.
Habitat Preferences
The preferred habitats of Milvago, Phalcoboenus, and Polyborus
include a variety of nonforest types: open scrub, grassy pampas,
tussock-grassland, pastures, treeless cultivated farmland, open
thorny chaco, savanna woodland, and open plantations. Phalco-
boenus albogularis may be an exception, since Olrog (1948: 478;
1950: 520) reports it as a forest bird.
Both species of Daptrius are inhabitants of tropical forest and
contrast markedly with the other caracaras in this feature. The
distribution of ater and americamis corresponds to the distribution
of wet lowland rain forest in Central and South America, but the
actual preference of these species seems to be less for the forest
interior than for more open situations within or along the forest:
river banks, small clearings, secondary growth, mangroves, and
tree-tops. The plumage pattern of both species of Daptrius, espe-
cially D. americamis, is most similar to that of adults of Phalco-
boenus carunculatus and P. megalopterus, which inhabit high
Andean steppes, an environment that differs in every respect from
that of Daptrius. This basic morphological similarity among taxa
of widely distinct ecological preferences seems to be more easily
understandable on an hypothesis of close relationship than on one
of convergent evolution. The fact that the species of Phalcoboenus
mentioned above have a very different immature plumage, while
those of Daptrius are far less dimorphic between adult and imma-
ture, might be a correlate of environmental differences between the
two genera. In the open habitats of the high Andes there is never
more than one species of caracara at any one locality (diversity
= 1 ) . Because congeneric competitors are absent, the sharp
dimorphism between adults and immatures of Phalcoboenus may
consequently reflect the results of slightly relaxed selective pres-
sures. In the tropical lowland forests, however, there are two
sympatric species of caracaras (diversity = 2), so that selection
through interspecific competition might possibly limit the range of
intraspecific variability, thus resulting in the evolution of similarity
(monomorphism) between immatures and adults of Daptrius. The
difference in size (character divergence) between the two species
12 BREVIORA No. 355
of Daptrius, which are sympatric, may be relevant here. Being so
different from each other (no overlap in range of wing lengths),
they should compete very little for food (see Schoener, 1 965 ) . The
interspecific difference in size, together with the lack of dimorphism
between age categories in Daptrius, may be roughly equivalent
ecologically to the striking dimorphism within high Andean Phal-
coboenus, which do not have any sympatric congeners. In Milvago,
where the two species are largely allopatric, one observes a con-
siderable overlap in wing lengths between the two, but one of the
species is conspicuously dimorphic (chimachima) , whereas the
other is not. Therefore, the situation in Milvago appears inter-
mediate to that in both Daptrius and Phalcoboenus. I beheve this
intermediacy is also found in the habitats occupied by Milvago
(such as open woodland, savannas) that are more or less inter-
mediate between lowland wet forest (occupied by Daptrius) and
barren high Andean steppes (occupied by Phalcoboenus).
Summary of Character A nalysis
If the characters discussed above are examined separately, the
variation among some of them shows the following possible group-
ing within the caracaras. (a) Polyborus is distinct in nostril shape
from the other genera, (b) Daptrius and Phalcoboenus are closer
to each other than to other genera since both have a strikingly
patterned black and white adult plumage, (c) The immature plum-
ages of Milvago, Phalcoboenus and Polyborus appear extremely
similar to one another, and differ, as a group, from those of Dap-
trius. If both adult and immature plumages are used, together with
habitat preferences, the two species of Daptrius appear to stand
out against most other species. First, the immature plumage of
Daptrius, when distinct from that of the adult (as in D. ater), is
quite different from the brownish, streaked or barred immature
plumage of all other species except Phalcoboenus australis. Sec-
ondly, the forest habitat of the species of Daptrius differs from the
nonforest habitats of the species in the other genera, with the pos-
sible exception of Phalcoboenus albogularis. (The remaining char-
acters [naked facial skin and size] seem of httle or no use in
estabUshing groups within the caracaras.)
I can only conclude from this analysis that all caracaras appear
to be closely interrelated, but that Daptrius is less similar to the
other three genera than these are to each other.
1970 SPECIATION IN THE CARACARAS 13
CLASSIFICATION
The splitting of the caracaras into four genera, endorsed by most
taxonomists, does not seem to reflect properly the close relation-
ships of these birds. Since the lumping of all caracaras into a single
genus may be going somewhat too far in the opposite direction, I
suggest here a third possibiUty, which is to put the caracaras in-
habiting nonforest habitats in a single genus (Polyborus, including
Milvago and Phalcoboemis) and to keep the forest caracaras in a
second genus (Daptrius). Further subdivisions within the non-
forest caracaras can be made by using species-groups and super-
species, which do not burden the nomenclature with additional
names (as pointed out by Cain, 1954), yet permit a finer hierarchy
between the genus and species levels.
I present below a classification outline of the caracaras, includ-
ing species-groups and superspecies. Each species-group represents
a former genus. The grouping of some species in a superspecies
(included in braces) is given here in anticipation of the discussion
on speciation in the next section of this paper. The sequence of
taxa in this list is arbitrary, and does not pretend to suggest that
some taxa are more primitive than others, since such decisions
would be guess work.
Genus Dap /m/5 Vieillot, 1816 (forest caracaras)
D. ater Vieillot, 1816
D. americanus (Boddaert, 1783)
Genus Polyborus Vieillot, 1816 (nonforest caracaras)
1 . chimachima species-group
P. chimachima Vieillot, 1816
P. chimango Vieillot, 1816
2. plancus species-group
P. plancus (Miller, Mil) (includes lutosus Ridgway,
1876, considered by some authors as a separate species)
australis superspecies
iP. australis (Gmelin, 1788)
\p. megalopterus (Meyen, 1834) (includes carunculatus
(Des Murs, 1853), and albogularis Gould, 1837, con-
sidered by some authors as two separate species)
14 BREVIORA No. 355
VARIATION AND SPECIATION IN DAPTRIUS
Of the two species of Daptrius, only D. americanus shows geo-
graphical variation. This variation seems to be a cline of diminish-
ing size, as measured by wing length, from Guatemala southward
through Central America to South America. According to Brown
and Amadon (1968), the populations from "southern Brazil" may
show an increase in wing length over those from farther north in
South America. This increase is in fact quite sharp, and is illus-
trated by Figure 2. The disjunction may reflect absence of gene
flow between birds from the Amazon Valley and those from the
uplands of Brazil. The birds Hving in the coastal forests of south-
ern Brazil and in the gallery forests of the Parana-Paraguay drain-
age system of south-central Brazil may be ecologically isolated
from birds living in forests of the Amazon Valley by parts of the
central Brazilian plateau, which are covered by extensive open
savannas and campos.
Any inferences that might be drawn about the possible evolu-
tionary history of this genus are prevented by the considerable
sympatry between ater and americanus (see map 76 in Brown and
Amadon, 1968).
320 330 340 350 360 370 360 390 400 4 10
I \ \ \ \ 1 \ 1 \ 1
Guatem,, Hond.Nicar. Cos, Rica
Ecuador. Peru
Brazil (Goias, Mato Grosso)
Figure 2. Geographical variation of wing length in Daptrius americanus.
Measurements are in miHimeters. Horizontal bars: range of measurements;
vertical bars: means.
1970 SPECIATION IN THE CARACARAS 15
VARIATION AND SPECIATION IN POLYBORUS
1. The chimachima Species-Group
The two species of this group, which constitute the former genus
Milvago, are largely allopatric. Polyborus chimachima occurs in
southern Central America (Costa Rica and Panama) and in South
America from Colombia and Venezuela in the north to northern
Argentina in the south, and P. chimango from northern Argentina
southward to southernmost South America. They are sympatric,
however, over a relatively broad zone, including Rio Grande do
Sul in southern Brazil, parts of northern Argentina, Uruguay, and
Paraguay, and southern Bolivia westward to the foothills of the
Andes.
Polyborus chimachima shows color and size variation. Tail and
culmen length appear to vary clinally, and to increase from south
to north, whereas the reverse seems to be true of wing length (see
Fig. 3).
Polyborus chimango shows geographical variation that seems to
conform to eco-geographical rules, since the southernmost birds
are the largest (Bergmann's rule), and the birds occurring along
the wet, forested Andean slopes are darker than those living in the
drier, grassy and shrubby plains of central Argentina (Gloger's
rule).
Geographical variation in P. chimango has been recognized tax-
onomically by the naming of three subspecies. Two of them, temu-
coensis (Andean slopes birds) and chimango (open plains birds)
intergrade broadly. The third subspecies, juegiensis, seems re-
stricted to the island of Tierra del Fuego, but is probably merely
the southward end of a north to south cline of increasing size.
Interestingly, the southern populations, from Tierra del Fuego and
the mainland of southern Patagonia, appear to leave their breeding
grounds to migrate northward as far as northern Argentina (Olrog,
1962: 112-113) in the southern hemisphere winter.
Although P. chimachima and P. chimango are undoubtedly very
close relatives, they have diverged morphologically rather consider-
ably. One of the differences has been mentioned earlier: the im-
mature plumage of chimango is very similar to that of the adults,
whereas the immature chimachima is quite unlike the adult. An-
other difference between the two species may be ecological. As I
16
BREVIORA
No. 355
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1970 SPECIATION IN THE CARACARAS 17
judge from personal observations and literature records, P. chima-
chima may generally prefer somewhat denser, more wooded, habi-
tats than P. chimango, but no study of a possible habitat segrega-
tion has been undertaken in the area of sympatry. If tarsus length
reflects whether a bird is more or less arboreal, then variation
between P. chimachima and P. chimango in this character is in-
structive. That P. chimachima populations have shorter tarsi than
those of P. chimango (see Fig. 3) suggests that the latter may be
less arboreal than the former. Whether this difference is true
character divergence should be examined carefully.
Brown and Amadon (1968: 739) say that the two species "may
be regarded as a super-species." Even if a broad superspecies con-
cept is embraced, it seems to me that the zone of overlap, con-
sidered together with the morphological differentiation between
chimachima and chimango, clearly suggests that these two taxa are
past the species borderline, and that application of the superspecies
concept to their case may no longer be correct. As far as I know,
hybridization does not take place between the two species where
they are sympatric; a field study of the pair in their overlap zone
would nevertheless be rewarding.
It seems most likely that the pair of species chimachima and
chimango has originated by a straightforward process of splitting
of one ancestral population into two. Yet secondary sympatry is
too extensive to permit more speculation about the original
isolation.
2. The planciis Species-Group
This group consists of P. plancus and the P. australis super-
species. The birds of this group are medium to large, and have
brownish immature plumage with dark streaking and/or barring.
The P. plancus species-group includes both the former genera
Polybonis and Phalcoboemis. Brown and Amadon (1968: 730)
remarked that "Phalcoboemis australis is in all respects inter-
mediate" between Polyborus and Phalcoboenus, yet they kept the
two genera distinct.
P. plancus is essentially a lowland species, while the members of
the P. australis superspecies occur mostly in the Andes, in some
localities at high altitudes. Where plancus meets members of the
australis superspecies, as on the Falkland Islands and on islands
off Tierra del Fuego, the two may live in habitat co-occupancy,
but their ecological relationships have not been studied. In any
18 BREVIORA No. 355
event, sympatry in the P. plancus species-group is very limited and
geographically peripheral.
Polyborus plancus
Polybonis plancus has a broad distribution from the southern
United States and Mexico to Tierra del Fuego and the Falkland
Islands and exhibits extensive geographical variation. The birds
from Florida, isolated from the remainder of the mainland North
American birds, are closer geographically and morphologically to
the Cuban ones (both are included in the subspecies auduboni) .
The birds of the population living on the Tres Marias Islands off
western Mexico (subspecies pallid us) are morphologically differ-
entiated from the nearest mainland populations (for a discussion
of the characters of palUdus, see Grant, 1965: 12-14). The con-
tinental population living from eastern Panama southward to
northwestern Peru near the Upper Maranon and Amazon Rivers
(subspecies cheriway) are well marked. There is intergradation
between cheriway and southern South American birds (subspecies
plancus) in Brazil (see Hellmayr and Conover, 1949: 283-284).
The extinct Polyborus lutosus lived on Guadalupe Island (see
e.g., Abbott, 1933; Greenway, 1958). The adults looked very
similar to plancus, but white was replaced by brown, and there
was no black on the abdomen; the immature was brown and
streaked. This insular population was certainly well marked; it
is considered here as having been a strong subspecies of plancus.
Brown and Amadon (1968: 736) maintained lutosus as a separate
species.
The australis Superspecies
The four nominal species (the former genus Phalcoboenus) in-
cluded in this superspecies are Andean and Patagonian in distribu-
tion (Fig. 4). One of the species, australis, occurs only on islands
off southern South America. The three others, carunculatus, mega-
lopterus, and albogidaris, occur along the Andean cordillera, from
Colombia to Tierra del Fuego.
Polyborus australis is larger than the three other species, but
resembles them in several other respects. The pattern of breast and
abdominal streaks in the adults is especially reminiscent of the
geographically distant carunculatus. P. australis breeds on the
Falklands, where, according to Cawkell and Hamilton (1961), its
numbers have decreased in the recent past. It also breeds on sev-
eral islands off Tierra del Fuego (Staten and Navarino), and on
islands of the Cape Horn Archipelago (for example, Grevy, Bayly,
1970
SPECIATION IN THE CARACARAS
19
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Figure 4. Geographical distribution of the Folyborus aiistralis superspecies.
A question mark represents the alleged occurrence of albogulans at Puente
del Inca, Mendoza (see Hellmayr and Conover, 1949: 217-218). Labeled
localities are identified in the text.
20 BREVIORA No. 355
Freycinet, Hershel: Olrog: 1950: 520). Reynolds (1932) ob-
served one P. australis on Woodcock Island in the Beagle Channel,
and Olrog (1948: 478) saw it at Yendegaia, along the southern
coast of Tierra del Fuego, where he also collected P. megalopterus
albogularis (see Fig. 4). This observation (not mentioned by
Brown and Amadon, 1968: 731 ) implies that P. australis and P.
megalopterus albogularis are narrowly sympatric on the main island
of Tierra del Fuego, but this suggestion needs careful checking in
the field. P. australis is sympatric with P. plancus on some islands
off Tierra del Fuego and the Falklands.
Polyborus carunculatus, megalopterus, and albogularis are here
treated as three subspecies of a single species, as previously advo-
cated by Hellmayr and Conover (1949: 276-277), and adopted
later by some students of South American birds (e.g., Olrog, 1963;
Koepcke, 1964; and Johnson, 1965). (The name megalopterus
Meyen, 1834, has priority over albogularis Gould, 1837, and
should be used as the specific name, contra Hellmayr and Conover,
1949.) Other authors, however, have maintained the three taxa
as separate species (e.g., Amadon, 1964: 14; de Schauensee, 1966;
Brown and Amadon, 1968).
The divergence of opinion about the taxonomic status of these
three taxa has hinged on two things: first, the striking interspecific
adult plumage color differences, and secondly, the allopatry of the
taxa. A third factor, namely the absence of intermediate specimens
between any two of these forms, has been invoked by some authors
to justify their treating the three as species. Brown and Amadon
(1968: 371), for instance, stated: "the two [taxa] that are the
most similar, albogularis and megalopterus, are not known to inter-
grade rather [sic] there seems to be a slight gap between their
ranges, suggesting friction." A few specimens, however, do seem
to be intermediate between carunculatus and tnegalopterus on the
one hand, and between megalopterus and albogularis on the other,
as discussed below.
The characters that vary from one to the other of the three taxa,
carunculatus, megalopterus and albogularis, are: the amount of
white at the tip of the primaries, the degree of curliness of the crest
feathers, the extent of bare skin on the throat, and the coloration
of the underparts. The two northern taxa, carunculatus and mega-
lopterus, live in high Andean grassland and scrub above the timber
line, at altitudes usually higher than 3000-3500 meters. Both of
these forms are common birds where they live, and can be seen
1970 SPECIATION IN THE CARACARAS 21
daily in small groups or singly (personal observation). The popu-
lations of carunculatus are geographically isolated from those of
we-^alopterus by a hiatus that includes the low Andes of northern
Peru, the depression of the Upper Maranon Valley and other val-
leys in northern Peru (see Fig. 4). Birds north of this gap {carun-
culatus) have the throat and breast black with white longitudinal
streaks or spots, and the abdomen white. South of the gap, birds
{megalopterus) have throat and breast black without white spotting
or streaking, and a white abdomen.
These morphological differences, considered in the light of the
ecological barrier lying between the two taxa, might be interpreted
as reflecting a period of geographical isolation during which differ-
ential selection acted on populations cut off from free gene flow.
Absence of gene flow, however, seems surprising in view of the
good flying abilities and the relative abundance of birds of this
group. It is therefore noteworthy that Zimmer (1930: 248) found
three adult males of megalopterus from Peru, all showing "an
interesting progression in the direction of carunculatus." Two of
the three specimens (from Macate, Department Ancash, and
Panao, Department Huanuco, see Fig. 4), have small, pale or
whitish spots on the lower breast; the third specimen (from Junin,
farther south) lacks breast spots. Zimmer's description of the three
birds should be consulted for additional details.
These data seem to indicate that gene flow is indeed possible
between Ecuadorian carunculatus and Peruvian megalopterus.
Further intensive collecting of these birds in northern Peru is badly
needed. The total number of specimens of either carunculatus or
megalopterus from areas close to the barrier of northern Peru is
low, so hybrid specimens might appear to be much rarer than they
really are.
What about the situation between megalopterus and albogularis?
The northern megalopterus is a common bird in the high Andean
scrub vegetation of the Argentine-Chilean cordilleras. The south-
ern albogularis seems to occur in Nothojagus forests rather than
open vegetation types (see Olrog, 1948: 478; 1950: 520; Philippi
et al., 1954: 39). There seems to be a distributional hiatus (see
Fig. 4) between the northernmost records of albogularis (in
Neuquen, fide Olrog, 1963: 116) and the southernmost ones of
megalopterus (in Talca, fide Johnson, 1965: 265). We do not
know, however, whether this gap is real or not, because of the
general scarcity of collections made in the "hiatus" area. In any
22 BREvioRA No. 355
event, what is real is the fact that no ecological barrier interrupts
the distribution of megalopterus and that of alhogularis in the way
the northern Peruvian low does between megalopterus and carun-
culatus. The situation seems therefore more complex ecologically,
and deserves field study.
From these considerations, contact should be possible between
megalopterus and alhogularis, and if they are not reproductively
isolated, gene flow should occur between them. In February, 1965,
at 1950 m on Cerro Catedral, near Nahuel Huapi, Rio Negro,
Argentina (see Fig. 4), I observed two adult Polyborus that were
attracted by a small garbage dump near the Refugio Lynch. One
of them had the throat and breast black, in contrast with the white
of the abdomen (megalopterus-likQ phenotype). The second had
both a white throat and breast, with only the sides of the breast
black, not forming a black pectoral band (albogularis-YikQ pheno-
type). This observation might have been of a mixed pair, but
unfortunately the birds could not be collected.
The possibility of mixed pairs and of offspring from them, seems
to be shown by two adult birds that exhibit what appears to be
intermediacy between megalopterus and alhogularis. One of these
birds, an adult male taken in February, 1 896, in Chubut, southern
Argentina, was described by Scott (1910) as Ihycter circumcinctus.
The underparts of this specimen are as follows: the throat is white,
followed by "a band of black below the throat patch more or less
variegated by white on some of the feathers; this band is about an
inch in width; lower part of the under neck pure white, the black
of the sides of the neck confining the white of this region to a
narrow area, widening into the pure white of the breast." This bird
appears to be like a specimen of alhogularis with a narrow black
breast band. The second of these birds, an adult male from Nahuel
Huapi (see Fig. 4), is mentioned by Hellmayr and Conover (1949:
277). This specimen (British Museum 99.1.27.229) has a distinct,
interrupted breastband.
The same conclusion seems, therefore, to follow from these
scanty data as from those on carunculatus and megalopterus: con-
tact between megalopterus and alhogularis seems a reality since at
least two specimens are somewhat intermediate between the two
taxa, and since birds from the two phenotypes were sighted to-
gether. Amadon, however, speaking of Scott's "circumcinctus,"
said that it "may be a mutant rather than a true genetic intergrade"
(1964: 15). This hypothesis seems unlikely to me. Since speci-
mens of alhogularis are relatively rare in museums, the two birds
1970 SPECIATION IN THE CARACARAS 23
with a black pectoral band represent a relatively high frequency of
the black-banded phenotype, perhaps as many as 1 in 10 or 1 in
15 — a number too high to be accounted for solely on the basis of
recurrent mutation. A third possible interpretation would be that
there is a polymorphism involving breast color. If this were true,
then obviously megalopterus and albogidaris should be considered
members of the same species.
For the time being, I believe the best interpretation of the situa-
tion in this complex to be that the three taxa, although clearly dif-
ferentiated morphologically as adults, have not achieved compbte
reproductive isolation, so that when two of them come in contact,
whether across a barrier (carunculatus with megalopterus) or not
(megalopterus and albogularis) , they produce hybrids. The tax-
onomic solution I propose in this paper is to consider all three
taxa conspecific, although I realize that this lumping may be a
little premature in view of the paucity of data. If additional study
should reveal that hybridization is very limited, even though there
may be plenty of opportunity for it to take place, then it would be
justified to maintain the three taxa as species, although it would
be necessary to emphasize that they are really semispecies: a truly
intermediate stage in the speciation process.
DISCUSSION
The caracaras are interesting to the student of speciation, be-
cause they offer a variety of phenomena that are interpreted as
intermediate in the process of species formation (see Table 3).
They can be summarized as follows. In Daptrius americanus, the
populations from southern Brazil may be geographically isolated
from other populations farther north in South America. In Poly-
borus plancus can be seen phenomena of incipient spsciation.
Isolated populations, both insular (Tres Marias Islands, Cuba)
and continental, exist, showing varying degrees of morphological
differentiation. The most differentiated population (lutosus) was
wholly insular, yet was of no evolutionary significance for further
speciation since it is now extinct. The other insular populations,
on the Tres Marias Islands and Cuba, are much less differentiated
than lutosus, the Cuban one even being similar morphologically to
the Florida population. On the continent, populations from north-
em South America (cheriway) are sufficiently differentiated from
southern South American ones (plancus) for some ornithologists
24
BREVioRA No. 355
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1970 SPECIATION IN THE CARACARAS 25
to have treated them as species. There is some evidence that (sec-
ondary? ) hybridization takes place near the mouth of the Amazon
where they come into contact.
The taxa of the Polyborus australis superspecies present another
"stage" of the speciation process, in that one of the members of
the superspecies, australis, is sufficiently distinct to be considered
unhesitatingly as a species. The three remaining members, how-
ever, present interesting situations. In one instance, differentiation
seems to have taken place across an ecological barrier (the low
area of the northern Peruvian Andes), yet hybridization appears
to occur in spite of this gap. In the second instance, no barrier is
evident today, and some hybridization seems to occur. The south-
ernmost taxon of this complex {albogularis) apparendy meets
australis in Tierra del Fuego.
In the Polyborus chimachima species-group, speciation is com-
pleted, and the two species overlap now over a considerable area,
although they are allopatric over the major portions of their respec-
tive ranges. Finally, the two species of the genus Daptrius are so
different morphologically and show so much sympatry that recon-
struction of their history is impossible.
Although six of the seven species I recognize in the caracaras
have extensive geographical distributions, the existing patterns of
speciation, or incipient speciation, seem to indicate that multiphca-
tion of species has occurred mostly through the formation, and
subsequent differentiation, of small or relatively small peripheral
isolates. The present isolates of Daptrius americanus and Polyborus
plancus are restricted to small areas around the periphery of the
range of the species, and the geographical location of Polyborus
australis relative to Polyborus megalopterus seems to suggest for-
mer peripheral isolation of the first named species. Differentiation
within P. megalopterus does not seem to correspond as clearly to
a pattern of isolation in peripheral areas, although the central
populations {megalopterus) do have a much broader distribution
than either the northern (carunculatus) or southern (albogularis)
ones.
The previous summary of speciation in the caracaras shows that
this process is most actively taking place along the Andes, where
the various members of the Polyborus australis superspecies live.
In the other, lowland, taxa, the speciation process is either com-
pleted (as in Daptrius or the Polyborus chimachima species-group)
or is not as pronounced {Polyborus plancus, Daptrius americanus) .
26 BREVIORA No. 355
It is tempting to relate the apparently greater evolutionary activity
in the Andean caracaras to the recent geological history of this
Cordillera. The high Andean grasslands and scrub habitats (paramo
and puna) where Polyborus megalopterus now lives are undoubt-
edly the most recent environments of the Andes, and cannot be
older than the latest phases of uplift, which brought the mountains
to their present tremendous altitudes during the Pho-Pleistocene
(see Childs and Beebe, 1963; Steinmann, 1930; Ahlfeld and
Branisa, 1960; and Briiggen, 1950; for summaries of the geological
development of the Andes). The differentiation within P. megalop-
terus most probably occurred during the Pleistocene glaciations,
although to attempt the dating of such processes is almost complete
guess work. However, if we recall that during glacial episodes, the
temperature depression lowered the altitude of the upper vegeta-
tion zones, where P. megalopterus lives, then it becomes possible
to envision the separation of a northern isolate (proto-carunculatus)
in Ecuador at interglacial time, when the altitudinal raising of this
treeless zone occurred, thus increasing the effectiveness of a natural
barrier such as the Upper Maranon Valley and northern Peruvian
low for birds living on either side of it. The isolation of caruncu-
latus from megalopterus, or, rather, of proto-carunculatus from
proto-me galopterus, might, then, have happened during an inter-
glacial. It is, of course, not possible to suggest which of the several
interglacial periods was responsible for such an event.
Similar glacial-interglacial oscillations may have permitted the
separation of proto-megalopterus and proto-australis in extreme
southern South America. During the maximum glacial, extreme
southern South America was covered with an ice-sheet (Caldenius,
1932; Polanski, 1965) which probably forced Andean biota to
"retreat" considerably northward. At the same time, however, the
Falkland Islands were left unglaciated, and, furthermore, were of
greater area than today because of a concurrent lowering in sea-
level. It seems therefore possible that during the maximum glacia-
tion the southernmost populations of the stock common to mega-
lopterus and australis remained on a Falkland refuge, where they
were geographically isolated from mainland populations by the ice
barrier, added to the sea barrier. If such a separation did indeed
take place during the maximum glaciation, which is attributed to
the late Pleistocene (Wiirm or Wisconsin) (see Polanski, 1965),
then the splitting of an ancestral stock into the modern australis
(having evolved from a population in a southern insular refuge)
1970 SPECIATION IN THE CARACARAS 27
and megalopterus (having remained in Andean Patagonia, but con-
siderably farther north than its present-day southernmost limit)
may have taken place as recently as 50,000 to 80,000 years ago.
Of course, it is also possible that the original separation took place
during an earlier, somewhat less extensive, glacial episode, but
since the maximum glaciation apparently obliterated earlier re-
mains, it is futile to speculate any further about the possible course
of this event.
ACKNOWLEDGMENTS
I thank Ernst Mayr for his continued advice throughout my
studies of speciation in Andean birds. He and Dean Amadon
criticized an earlier version of this paper. W. John Smith and
William H. Drury, Jr. kindly allowed me to use their field notes
on several species.
Field work was financed by the Frank M. Chapman Memorial
Fund of the American Museum of Natural History, the National
Science Foundation (grants G-19729 and GB-3167 to the Com-
mittee on Evolutionary Biology of Harvard University), the Society
of the Sigma Xi, and the Bourse federate de voyages of the Societe
helvetique des sciences naturelles (Switzerland).
I am grateful to the following persons, who helped me in various
ways during my examination of specimens: Jean Dorst (Museum
national d'histoire naturelle, Paris), I. C. J. Galbraith (British
Museum, Natural History), Father Antonio Olivares (Instituto de
Ciencias Naturales, Bogota), Raymond A. Paynter, Jr. (Museum
of Comparative Zoology), and the late R. A. Philippi (Museo
Nacional, Santiago). Finally, I acknowledge the assistance of
Warren Hubley during the course of this study.
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1970 SPECIATION IN THE CARACARAS 29
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BREVIORA
Mnaseiiamn of Coimparative Zoology
Cambridge, Mass, 30 November, 1970 Number 356
On new species in a new earthworm genus
from Puerto Rico^
G. E. Gates2
Abstract. A new genus of earthworms, Estherella (family Glossosco-
lecidae), with new species montana (type species) and nemoralis, is
described from Puerto Rico, and its remarkable structural modification is
discussed.
A seemingly impossible evolutionary modification was shown
by several worms received 25-30 years ago. Immaturity, amputa-
tion, maceration, and paucity of specimens prevented completion
of species descriptions to desired standards. Unfortunately subse-
quent material never became available. Various attempts to secure
it were futile. This contribution now is proferred in hope of
awakening interest in: 1) An evolutionary development that some
zoologists categorically maintain is impossible. 2 ) The mostly
unknown earthworm faunas of Caribbean Islands.
Glossoscolecidae
Estherella gen. nov.
Definition. Digestive system, with a gizzard in iii, three pairs of
calciferous glands in v-vii, each gland sausage-shaped, vertically
placed alongside gut, with a short duct from dorsal end opening
into esophagus just lateral to the supra-esophageal vessel, in-
testinal origin in region of xix-xx, with a lamelliform typhlosole,
without caeca and supra-intestinal glands. Vascular system, with
a single dorsal trunk aborted in front of hearts of iv, complete
ventral and subneural trunks, the latter adherent to parietes, a
supra-esophageal trunk in v-xiv, paired latero-esophageal trunks
1 From research financed by the National Science Foundation.
2 Zoology Department, University of Maine, Orono.
2 BREVIORA No. 356
in iii-ix with connectives to supra-esophageal in v-vii. Hearts, in
iv-vii lateral, in viii latero-esophageal. Nephridia, holoic and
vesiculate. Nephropores, obvious, in a regular longitudinal rank
on each side in region of CD. Pigment, none. Septa, present from
3/4. Prostomium, none, replaced by a protrusible proboscis. Setae,
eight per segment, in regular longitudinal ranks.
Quadrithecal, spermathecae adiverticulate, pores in region of
CD, at 5/6-6/7.
Type species, E. montana n. sp.
Distribution. Puerto Rico.
Estherella montana sp. nov.
Puerto Rico, El Yunque Mountain. Wet cloud forest at
± 2500 feet. May 1938. 3-0-0. P. J. Darlington per G. E.
Pickford. (Mus. Comp. Zool.) About 2200 feet, an anterior
fragment, C. W. Richmond & L. Stejneger. (U. S. Nad. Mus.)
External characteristics. Length, 50-60 mm (juveniles), 175
mm (posterior amputee of 148 segments). Diameter, 4 mm
(juveniles), 12 mm (amputee). Segments of 54 mm juvenile,
188. Color, white (long alcoholic preservation). Peristomium,
much shorter than ii but of about the same appearance externally
as subsequent segments. Intersegmental furrows, distinct. Seg-
mental length, gradually increasing posteriorly to region of vii-ix.
Secondary annulation, a presetal and a postsetal secondary furrow
in each of ix-xxvi, postsetal secondaries unrecognized behind
xxvi. Nephropores, obvious, present from ii, well behind inter-
segmental furrows but usually less than half way toward segmental
equators, at or near C. Setae, paired, ventral couples first certainly
recognizable in v, lateral couples in region of xv, in region of xxx
CD slightly < AB much < AA < BC, posteriorly A A ca. - BC,
still further back AA ^ BC. Dorsal pores, none.
Quadrithecal, spermathecal pores, minute, superficial, in CD,
at 5/6-6/7. Other genital apertures, unrecognizable. Genital
tumescences, transversely and shortly elliptical, indistinctly de-
limited, each with two circular areas of epidermal translucence at
center of which is a follicle aperture, a,b/xw-\xi\. Region ot AA,
rather deeply depressed through xv-xxiv.
Internal anatomy. Septa, 4/5-13/14 thickly muscular to
muscular, funnel-shaped, large, apices well posteriorly, 14/15 and
following septa slightly strengthened by muscular fibers. Septum
3/4, a delicate transparent membrane bearing on its posterior face
1970 NEW EARTHWORM GENUS 3
one pair of nephridia and on its anterior face two other pairs of
tubules, with insertion on gut immediately behind gizzard. Pig-
ment, if once present in body wall, completely leached by pre-
servative. A large, empty canal, ellipitical in cross section,
apparently completely circumferential, in anterior portion of body
wall in each of i-x. Canal size, decreasing posteriorly. Brain, in
ii. Nerve cord sheath, massively muscularized (Fig. 1) anteriorly
but so as to leave a greyish translucent line visible at mD and
mV in the cord between segmental ganglia (Fig. 2).
Buccal cavity, in i dorsally of small juveniles, seemingly pro-
vided, though only temporarily, with a suckerlike pad somewhat
resembling the withdrawn and depressed condition of the prosto-
mium in various megadriles, in ii dorsally with a circular aperture
into a tunnel containing a presumably protrusible proboscis 1-2
mm long. Gut from level of intersegmental furrow 2/3 to septum
4/5, ca. 30 mm long, sigmoid, comprising a pharynx (4 mm
long), a bulb (5 mm thick dorsoventrally and with a glandular
chamber anteriorly), a slender esophagus (14+ mm long) with
closely crowded, low longitudinal ridges on its inner wall, a sort
of conical crop (5 mm long) with circular ridges on its inner wail,
and a powerful gizzard (6 mm long) referable to iii.
Calciferous glands, in contact with each other mesially under
the gut, in vii ducts longer but concealed by adherence of 7/8
(near apex of its funnel) to the gut. Typhlosole, present from
region of xxiv-xxvi, 10 mm high (3 mm, small juvenile), rolled
up on itself like a scroll, ending in region of 125th segment (132d
of 188). Lateral typhlosoles, not lamelliform, rounded and pro-
tuberant ridges in first one or two typhlosolar segments.
Ventral blood vessel, high up in coelom and near gut in v-xi at
least. Extra-esophageals. interconnected by a transverse vessel
just in front of 4/5 and just under the ventral trunk, anteriorly
passing up and branching among nephridia associated with 3/4.
Supra-esophageal, with a large branch on each side in v and vi
that bifurcates, one branch to a calciferous gland near the duct,
the other passing down along anterolateral aspect of the gland to
an extra-esophageal trunk, connected also with extra-esophageals
by a pair of vessels, seemingly on posterior face of 7/8 but mostly
within the septum. Hearts, of iv-vii slender and lateral, of viii
apparently latero-esophageal — posterior bifurcations to dorsal
trunk slender and empty, anterior branches filled with blood and
obviously joining the supra-esophageal.
4 BREVIORA No. 356
Nephridial ducts (of anterior segments), passing down through
longitudinal muscle layer and then turning forward to cross the
circumferential intra-parietal canal, thence anteriorly widened and
with more opaque (muscularized?) wall.
Spermathecae, rudimentary, adiverticulate, within the longi-
tudinal muscle layer. ,
Remarks. Small juveniles are assumed to be of the same species
as the large worm from the same mountain.
Rudimentary state of the spermathecae and absence of macro-
scopically recognizable gonads, funnels, and seminal vesicles, in-
dicate that even the large worm was juvenile though maximum
diameter for the species may have been attained.
The parietal insertion of the delicate septum 3/4 had become
unrecognizable presumably as a result of pinning out the specimen
after a longitudinal incision had been made near the mid-dorsal
line.
Any connection between nephridial ducts and the circumferential
intraparietal canals would have been too small to recognize in
dissection, and microtome sections were unsatisfactory. The
canals were crossed diagonally by delicate fibers (or septa?).
Similar canals had been observed at least once before, but records
were destroyed during World War II.
Abortion of dorsal trunk in front of hearts of iv and posteriorly
in that segment was found in each dissected specimen considered
herein. That and other characters already mentioned in the generic
definition do not need mention again in species descriptions.
Photographs of nerve cord sections were provided by Prof. E.
Carpenter.
E. nemoralis sp. nov.
Puerto Rico. Luquillo Forest (Caribbean National Forest),
La Mina Recreational Area, at 1800 feet, February 22, 1947.,
2 macerated specimens (several younger specimens possibly of
the same species, also macerated.) R. Kenk. (U. S. Natl. Mus.)
External characteristics. Length, 250 mm. Diameter, 9 mm.
Segments, 220 (at 195/196 a tail regenerate with terminal anus).
Color, possibly red originally, even after long alcoholic preserva-
tion dorsum with a slight reddish tinge, except in regenerate.
Nephropores, obvious, present from ii, in CD. Setae, closely
paired throughout, AB = CD, AA > BC, DD ca. = VzC, ventral
1970 NEW EARTHWORM GENUS 5
couples of some segments modified (? but genital tumescences not
recognized).
Clitellum, perhaps represented by a dark brown coloration in
xv-xxii, xxiii/eq, which is conspicuously lacking in a small area
around each nephropore, no epidermal tumescence recognizable.
Ouadrithecal, pores minute, superficial, each at center of a small
tubercle at C and at or immediately in front of 5/6-6/7. Female
pores, postsetal in AB of xii(?).
Internal anatomy. Septa, 4/5-6/7 very thickly muscular, a
transparent, funnel-shaped membrane bearing three pairs of
nephridia inserted on the gut just behind the gizzard almost im-
mediately in front of 4/5 presumably being 3/4, 7/8 lacking or
else inserted on parietes over intersegmental furrow 8/9.
Calciferous glands, without a central lumen but with a honey-
comb appearance in cross sections, each with a small distal
appendage.
Supra-esophageal, bifurcating posteriorly in xiv, giving off two
pairs of vessels, one immediately behind the other and both just
in front of the septum in each of v-vii, the posterior of each pair
giving ofl" branches to the calciferous gland and ventrally joining
the extra-esophageal trunk of it^ side, the anterior vessels passing
onto stalks of calciferous glands and down through the glands into
the terminal appendages. Extra-esophageal, first visible in region
of 3/4 as a result of union of several large vessels, with several
branches to each calciferous gland of its side. Subneural trunk,
large, zigzag-looped, closed ends of loops visible beyond both sides
of the nerve cord, bifurcating just in front of subpharyngeal gang-
lion (one specimen) or in region of xvii (one), each branch
passing anteriorly in a zigzagged course lateral to the cord but
connected with its twin on the opposite side by numerous trans-
verse vessels. Ventral trunk, high up in coelom as in E. montana.
Hearts, large, two pairs, possibly latero-esophageal and attributable
to viii-ix (?).
Testis sac (or sacs?) filled with coagulum, surrounding or
including hearts belonging in viii (?).
Spermathecae, sessile, ducts confined to body wall, ampullae
small, protruding only slightly into coelomic cavities of vi and vii.
Remarks. Intersegmental furrows, in spite of the maceration,
are distinct, and septa 4/5-6/7 are inserted on the parietes directly
over intersegmental furrows 4/5-6/7.
Thickness of the subneural trunk is greater than that of the nerve
cord even in regions of segmental ganglia.
6 BREVIORA No. 356
The cuticle was loose and setae had been pulled out of their
follicles.
Repetition in the description of characters shared identically
with E. montana seems unnecessary. Mention should be made of
the fact that a proboscis and circumferential parietal canals were
not seen.
This species is distinguished from E. montana by absence of the
marked muscularity in the nerve cord sheath.
Estherella sp.
Puerto Rico. Luquillo Forest (Caribbean National Forest),
La Mina Recreational Area, 1800 feet, February 22, 1947, 1
macerated specimen. R. Kenk. (U. S. Natl. Mus.)
External characteristics. Size, 150 by 7 mm. Nephropores,
obvious, present from ii, in CD. Setae, paired throughout (ventral
couples modified in some clitellar segments?).
Qitellum, xv-xxii (and xxiii?). Tubercula pubertatis, longi-
tudinal bands of translucence, just lateral to B, each demarcated
laterally by a deep furrow. Female pores, postsetal in /4B of xii(?).
Internal anatomy. Male funnels, one pair, iridescent, imbedded
in coagulum apparently also containing hearts of viii and possibly
in sacs (or a testis sac?). Spermathecal ampullae, spheroidal,
without spermatozoal iridescence, slightly protuberant into
coelomic cavities from the angles of septal insertions and parietes.
Remarks. Setae had been pulled out of their follicles as in the
types of nemoralis. Gonads and female funnels were not found.
Structure, so far as could be determined, is the same as is shared
by the two preceding species.
The reason for anticipating a third species is the maturity at a
size smaller than is expected for the other two.
SYSTEMATICS
Cephalization has had little attention from oligochaetologists
and perhaps least of all in connection with a family in which one
manner of evolutionary modification seemingly had its most ex-
tensive, as well as perhaps least appreciated development.
Metamerism in oligochaetes sometimes has been said to be
homonomous, i.e., similar throughout the body. Typically, the
soma is in anteroposterior segments, each of which, at least in
earlier stages of evolution, has four pairs of setae, a pair of
1970 NEW EARTHWORM GENUS 7
nephridia. and a section of the gut. Such a segment is demarcated
externally from each of its two contiguous neighbors by inter-
segmental furrows, circumferential lines where the epidermis is
thinnest. Internally, a segment is deUmited by transverse parti-
lions, the intersegmental septa. The latter, typically, are in exactly
the same anteroposterior levels as the intersegmental furrows.
Peristomium and periproct, according to such definitions, are not
segments, though for practical purposes are counted as such.
Differentiation in a relatively short anterior portion of the soma
of special digestive organs such as gizzards, calciferous glands,
etc., and localization of gonads are aspects of oligochaete cephali-
zation so universal as not even to have been thought to be involved.
More usually considered were abortions, as of follicle and nephri-
dial anlage, and disappearance of septa and intersegmental fur-
rows, i.e., those structures existence of which enables recognition
of segments. Slight (or at least seeming) diplacements of septal
insertions on the parietes have been known for some time. Recog-
nized but recently (Gates, 1943: 92) was a seemingly posterior
dislocation of the parietal insertion of septum 9/10, in Pontoscolex
corethrurus (Miiller, 1857), to a position over site of inter-
segmental furrow 10/11. Failure to recognize that dislocation was
responsible in part for assignment of certain organs in systematic
descriptions to wrong segments. How many other errors of that
sort were made in past characterizations of glossoscolecids remains
to be learned.
Appearance of deep, secondary, and even tertiary furrows hardly
distinguishable from the primary intersegmentals, in association
with abortions of setae and nephridia, also has been responsible
for errors in determination of organ locations. Such mistakes are
unfortunate, as knowledge of exact organ position along the ante-
roposterior axis is of first importance for megadrile systematics
and phylogeny. Absence of data as to segmental location of the
gonads in the Puerto Rico species is not so regrettable as would
formerly have been thought since it is now known that "andry"
often, and sometimes even "gyny," does vary intragenerically.
Fortunately, the worms now under consideration, in spite of
the poor condition, are free of external modifications that might
lead to wrong determinations of organ locations. Sctal follicles,
to be sure, have been aborted in some of the anteriormost seg-
ments, but compensation is provided by the nephropores which
not only are all present but also are obvious. Secondary and
8 BREVIORA No. 356
tertiary furrowing is lacking, and the anterior segments are clearly
demarcated by unmistakable intersegmental furrows. The first
two segments have the normal appearance of externally exposed
epidermis. No evidence was found for the existence of an actual
pre-oral vestibule comprising one or two rudimentary segments
no longer exposed regularly to the external environment. Further-
more, the unusual condition next to be discussed characterizes
every individual of two, or possibly even three, species. Individual
abnormality or defective anterior regeneration accordingly cannot
be invoked as was contended when a single Panama specimen was
described (Gates, 1968) with a similar condition.
The powerful gizzard, being in front of a membrane that, be-
cause of nephridial relationships, must be regarded as septum 3/4,
is unusually anterior. With the single exception of the above-
mentioned Panama species, a megadrile gizzard has not hitherto
been found in front of segment v and often is further back in the
esophagus. Other organs also are too far forward. The last pair
of hearts is in viii, whereas the first pair of real hearts usually is
in ix. Calciferous glands are present in v only in the Panamanian
Thamnodriloides yimkeri Gates, 1968. Lastly, testes are at least two
segments in front of where they would normally be expected.
Accordingly, much more is involved than forward displacement
(homoeosis) of a single organ but rather a condition in which
all organs of the cephalic region from gizzard posteriorly are three
segments in front of their expected positions (regional homoeosis).
An initial or very early stage in an evolutionary development
that may have reached its climax in the Puerto Rico and Panama
worms is provided by P. corethrurus. In that species an inter-
segmental furrow between the first and second segments has di-
appeared along with the prostomium. The now rather flaccid
fusion metamere is small. Proof of what happened is provided in
many specimens by retention of the setae belonging to ii which are
now near the first intersegmental furrow that morphologically is
2/3. With loss of those setae, as in some individuals of the species,
the gizzard would have to be referred to v instead of vi. Organs
behind the gizzard also would be one segment anterior to their
usual position. Abortion of two further segments by the same
process under way in P. corethrurus would provide the regional
homoeoses of the Puerto Rico and Panama genera. Although
body wall and associated nephridia were markedly reduced or
deleted, the digestive system was not correspondingly shortened.
1970 NEW EARTHWORM GENUS 9
On the contrary, there has been so much elongation as to require
very considerable enlargement of associated septa into posteriorly
directed funnel-shapes. What happened in the nervous system may
prove to be interesting also.
Although homoesoses are identical, the digestive systems and
especially structure of the calciferous glands show that the two
fore-shortened genera are not closely related. Puerto Rican worms
may have evolved from a stock with calciferous glands in viii-x.
Genera so characterized are unknown. Related forms should be
sought to the south as the glossoscolecids evolved in tropical South
America. Puerto Rico now appears to be the northern limit of
generic endemism.
Completion of a development somewhat hke that now under
way in P. corethrurus may be responsible for attribution of testes
in Thamnodriliis matapi Righi, 1969, to segments ix and x instead
of the expected x and xi.
REFERENCES
Gates, G. E. 1943. On some American and Oriental earthworms.
Ohio Jour. Sci., 43: 97-116.
. 1968. On a glossoscolecid earthworm from Panama and
its genus. Megadrilogica, 1: 1-15.
10
BREVIORA
No. 356
1970 NEW EARTHWORM GENUS 11
LIST OF ILLUSTRATIONS
Figure 1. E. montana. Transverse section of nerve cord anteriorly to
show muscularization of the sheath.
Figure 2. E. montana. Transverse section of nerve cord anteriorly to
show the condition responsible for the appearance of a greyish translucent
line at mD and mV.
Figure 3. E. montana. Transverse section of nerve cord anteriorly to
show giant cells ventrally.
BREVIORA
MiLiiseiiinn of Coimpsirsitive Zoology
Cambridge, Mass. 30 November, 1970 Number 357
A review of the fossil Pelomedusidae (Testudines,
Pleurodira) of Asia
Roger Conant Wood
Abstract. The taxonomic status of the three Asiatic chelonian genera
that have been described as pelomedusids is reviewed. Of these, "Podoc-
neinis" imiica, although possibly a member of the family, is so poorly known
that familial assignment is not presently possible. Carteremys leithii and
Shwehoemys pilgrimi both appear to be valid palomedusid species. On
the basis of new material S. pilgrimi is redescribed and, in addition, a new
species of this genus from the Miocene of Baluchistan, S. gaffneyi, is
proposed.
INTRODUCTION
Living pelomedusid turtles are restricted to sub-Saharan Africa,
Madagascar, and South America. But paleontological evidence
indicates that pelomedusids formerly had a much more cosmo-
politan distribution; fossil representatives of this family occur in
North and South America, Europe, Africa, and Asia.
The purpose of the present paper is to review the extinct Asiatic
chelonian genera that have been described as pelomedusids as
well as to put on record two new fossil pelomedusid skulls from
Asia, one referable to Shwehoemys pilgrimi and the other repre-
senting a new species of the same genus.
Abbreviations used in this paper are:
ANSP — Academy of Natural Sciences, Philadelphia
BMNH — British Museum (Natural History)
GSI — Geological Survey of India
MCZ — Museum of Comparative Zoology, Harvard University
I have not seen the material described by Lydekker, Swinton,
and Williams that is contained in the collections of the Geological
Survey of India, but it is for the most part well figured in the
literature. Photographs of the type of Shwehoemys pilgrimi
Swinton have been available.
2 BREVIORA No. 357
PREVIOUSLY DESCRIBED ASIATIC PELOMEDUSIDS
Very few fossil pelomedusids have been described from any-
where in Asia, and none of these is particularly well known on the
basis of published material.
"Podocnemis*"' indica Lydekker 1887. This species was de-
scribed on the basis of a fairly complete carapace and on some
miscellaneous plastral fragments found at Nila in the Salt Range
of West Pakistan. Lydekker was uncertain about the age of "P".
indica but concluded (1887: 59) that it was probably of ". . . low-
est eocene . . . and may . . . correspond to the Cemaysian stage
of Reims, and the Puerco group of the United States." Since it
is now generally recognized that the Cernay beds are of late
Paleocene age and the Puerco beds of early Paleocene age, "P."
indica may actually be of Paleocene rather than Eocene age. Both
Paleocene and early Eocene deposits occur in the Tertiary sequence
of the Salt Range (Krishnan, 1960: 494), and too much uncer-
tainty exists as to the exact stratigraphic horizon from which the
only known specimen of "P." indica was obtained to resolve the
question of its age beyond all doubt. Invertebrates found in
association with the two chelonians {"Podocnemis" and Hemi-
chelys) described from this locality ". . . indicate that the bed
in which they were found is either of marine or estuarine origin;
and this is confirmed by the chelonians, one of which is covered
with the 'spaf of oysters, while sharks' teeth are embedded in the
matrix of the other" (Lydekker, 1887; 59-60).
Preservation of the carapace of "P." indica is rather poor;
Lydekker noted (1887: 60) that ". . . both specimens [from
Nila] . . . were in a much broken condition, and . . . required all
[the preparator's] skill to render them fit for description."
Lydekker's restoration of the carapace (1887, plate 13) indicates
that its most distinctive feature is a peak along the midline of the
neurals, with the pleurals on either side apparently sloping away
flatly rather than on a curve. The nuchal bone appears to be
unusual in that its postero-lateral sides are considerably longer
than its antero-lateral ones. The nuchal is rather small and does
not transgress the lateral boundaries of the first vertebral scute.
There are seven neurals; the first five are all longer than broad,
whereas the last two are broader than long. Two pleurals (the
seventh and eighth) meet in the midline between the last neural
and the suprapygal. No indentation occurs at the midline along
1970 ASIATIC FOSSIL PELOMEDUSIDAE 3
the anterior margin of the carapace and a cervical scute^ is lacking.
Whereas the first vertebral is nearly twice as wide as it is long,
the second and third vertebrals are both longer than broad.
Lydekker estimated (1887: 63) that the overall length of the
carapace must have been approximately 35 inches (87 centi-
meters), exceptionally large for a fossil pelomedusid. Critical
taxonomic evidence, such as whether or not the pelvis was fused to
the shell, the presence or absence of mesoplastra, and the scute
pattern on the anterior plastral lobe, is not preserved. Conse-
quently, there is no assurance that "P." indica is actually a pelo-
medusid, let alone a member of the genus Podocnemis. Neverthe-
less, its midline ridge and straight loping sides are somewhat remi-
niscent of the carapace structure of a recently discovered shell of
Shweboemys from Egypt (see p. 00), and the number, shape, and
arrangement of its neurals is typical of many pelomedusids, as is
its lack of a cervical scute. Therefore, it does not seem altogether
unlikely that "P." indica may represent the remains of some kind
of pelomedusid, although it probably is not a member of the genus
Podocnemis. Until more complete specimens of this taxon are
discovered, I do not think that a better identification of this speci-
men is possible than Pelomedusidae? incertae sedis.
Cartereniys leithii (Carter 1852). A second Asiatic pelome-
dusid taxon, "Hydraspis" leithii, was recovered from Intertrappean
beds near Bombay, India. Some uncertainty exists as to the age of
these sediments. Lydekker (1887: 60) regarded them as "lower
eocene." Referring to the age of the volcanics within which the
Intertrappean beds occur, Wadia (1953: 302) stated that "... it
is quite apparent that the Deccan Traps cannot be older than the
Danian stage of the uppermost Cretaceous [now Paleocene], while
. . . they cannot be much younger than the Eocene." According
to Krishnan (1960: 483-486), paleontological evidence afforded
by fossils contained within the Intertrappean beds is not par-
ticularly helpful for dating, but on other grounds he concluded
that the Deccan Traps ranged in age from late Cretaceous to,
perhaps, Ohgocene. Robinson (1970: 245) has expressed similar
views, stating that the Intertrappeans are ". . . probably mainly early
ij have here adopted the term suggested by Zangerl (1969: 315) for
this particular scute to avoid the confusing redundancy arising from the
conventional procedure of referring to both it and the underlying bone as
the nuchal.
4 BREVIORA No. 357
Tertiary in age. . ." Since the Intertrappeans of the Bombay
region are confined to the upper part of the Deccan Traps (Pascoe,
1964: 1385), their attribution to the Eocene would not be unrea-
sonable. Krishnan (1960: 482), Pascoe (1964: 1385) and
Robinson (1970: 245) all agree that the Bombay Intertrappeans
were laid down in fresh water, Pascoe further suggesting (1964:
1386) that the depositional environment was a shallow marsh.
Carteremys leithii was a small species; the only two complete
carapaces for which measurements have been recorded are IVk
and 8 inches (18-20 centimeters) in length (Carter, 1852: 187;
Williams, 1953: 6). Carter's original description was based on
material that is now unfortunately lost (Williams, 1953: 2). Only
three other specimens that can be referred to this species with
reasonable confidence have subsequently been discovered
(Williams, 1953: 6, and plate 3). The total number of neurals
is uncertain, although there are at least five. Evidently no cervical
scute was present, and the first vertebral was much smaller than
the second. The outer surface of the shell is covered with fine
sculpturing. In several respects the plastron is quite distinctive:
it is relatively narrow, with a semicircular anterior lobe projecting
well forward of the front of the carapace; between the broadly
rounded xiphiplastral tips is a very shallow anal notch, not com-
parable in its shape to those of any pelomedusid with which I am
familiar; and the outlines of the pelvic scars on the xiphiplastron
are also unusual. In spite of Williams' suggestion (1953: 4) that
small, laterally placed mesoplastra, a characteristic pelomedusid
feature, may have been present, there is no conclusive evidence
bearing on this point. None of the three existing specimens (all
belonging to the collections of the Geological Survey of India) is
sufficiently complete to demonstrate the presence or absence of
these structures (WiUiams, 1953: 6). Trapeziform gulars were
widely separated by an extremely broad intergular scute that
extended posteriorly to the humero-pectoral sulcus and thus pre-
vented the humerals from meeting in the midline also. The most
notable feature of the skull is the extensive emargination of the
roof from behind, a condition typical of most pelomedusids. A
well-developed jugal-quadratojugal bar is present. Although the
extremity of the mandibular rostrum was broken off in the
material Carter described, it is clear from what was preserved
that there must have been a broad, robust symphysis at the mid-
line of the lower jaw.
1970 ASIATIC FOSSIL PELOMEDUSIDAE 5
Long regarded as a chelid, "Hydraspis" leithii was eventually
redescribed and designated as the type of a new pelomedusid genus
by Williams (1953: 3-4). A combination of several characters —
the absence of a cervical scute, the relative proportions of the first
and second vertebrals, the remarkably large intergular, and the
lack of a parieto-squamosal arch combined with the presence of
a jugal-quadratojugal bar — led Williams to conclude that this
species could not be a cheHd. These characters, together with the
inferred existence of small, laterally placed mesoplastra, per-
suaded him that Carteremys was in fact a pelomedusid. On the
basis of the data presented in Table 1, I agree with Williams'
conclusion. If we disregard for a moment the question of whether
or not mesoplastra were present in Carteremys, then we see that
for all the characters tabulated, this genus and pelomedusids agree.
In contrast, none of the other families share more than two
characters with Carteremys. This comparison strongly suggests
that Carteremys is a pelomedusid. Should it eventually be possible
to determine that Carteremys had mesoplastra, the evidence would
overwhelmingly favor this determination.
Lydekker (1890: 22-23, fig. 2) referred an epiplastral frag-
ment, probably from Intertrappean beds and therefore possibly
of Eocene age, to "Hydraspis" leithii. Differing from Carteremys
in size and in gular-intergular proportions, this specimen may well
represent an unknown taxon, but too little is known of it to permit
useful discussion.
Shweboemys pilgrimi Swinton 1939. The type, and until now
only known specimen, of this species is a partial skull of Pliocene
or Pleistocene age from Burma (Swinton, 1939). Swinton be-
lieved that nasal bones, although not preserved on the specimen
he described, must have been present originally. Were this sup-
position true, his placement of the genus in the Pelomedusidae
would be suspect because one of the diagnostic characters of the
family is the absence of nasals (cf. Romer, 1956: 515). Evidently
Swinton was either unaware of the significance of this character
or else he did not consider it to be of great importance; at any
rate, he did not discuss its bearing on taxonomy. Nevertheless,
other observations led him to conclude (1939: 551) that
Slnveboetnys is a pelomedusid: 'in brief, the interest of the speci-
men is that in superior and lateral aspect there is little to dis-
tinguish it from the genus Podocnemis, while in palatal view it
has much similarity to Stereogenys. There is no doubt that it
BREVIORA No. 357
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1970 ASIATIC FOSSIL PELOMEDUSIDAE 7
differs from both genera and is a new form belonging to the family
Peiomedusidae."
NEW ASIATIC PELOMEDUSID MATERIAL
The recent recognition of a second, slightly more complete
specimen of Shweboemys pilgrimi by Mr. C. A. Walker in the
collections of the British Museum (Natural History) has provided
new information which confirms Swinton's familial determination.
Further supporting evidence is furnished by the new species of
Shweboemys from the Miocene of Baluchistan (West Pakistan)
described below. In addition, recent Yale University paleonto-
logical expeditions to the Fayum Depression of Egypt have pro-
duced conclusive evidence showing that "Podocnemis" antiqua
Andrews 1903 from the late Eocene Qasr el-Sagha Formation is
referable to Shweboemys. A detailed description of this species,
the only one represented by both skull and shell material, is being
prepared for publication elsewhere, but I include the diagnostic
characters of its shell in the emended diagnosis.
Shweboemys Swinton 1939
Type species. Shweboemys pilgrimi Swinton 1939.
Emended diagnosis. Skull with broad secondary palate formed
by medial expansion of maxillae and palatines, with narrow median
cleft extending posteriorly from behind premaxillae; outer border
of palatines not parallel to midline axis, but diverging from it at
an angle of approximately thirty degrees; laterally projecting,
prominent ectopterygoid processes; enlarged carotid charmels;
basisphenoid not covered by pterygoids ventrally. Carapace cordi-
form in outline, tapering to a point posteriorly; pleurals flat rather
than curved, sloping away from continuous midline ridge at gentle
angle; ventral surface of plastron completely flat; anterior lobe of
plastron very short and semicircular; posterior lobe narrower and
approximately one and one-half times longer than anterior;
outer margin of posterior lobe straight rather than curved, slanting
inward toward rear.
Referred species. Shweboemys antiqua (Andrews) 1903, S.
gaffneyi sp. no v.
Distribution. Late Eocene, Fayum Depression, Egypt; early
Miocene, Bugti HiUs, West Pakistan; Pliocene or Pleistocene,
Burma.
8 BREVIORA No. 357
The skull of Shweboemys is very similar to that of Stereogenys
but differs in several significant respects: the pterygoids do not
completely cover the basisphenoid ventrally; the lateral margins of
the palatines are not parallel to the midline axis; and the secondary
palate is less elongate, not extending back as far as the ectoptery-
goid processes.
Swinton's decision to base a new pelomedusid genus and species
on his Burmese specimen was founded on his belief that the skull
combined features of both Podocnetnis and Stereogenys without
being more like one than the other (see p. 5). That the palatal
structure of Shweboemys is very similar to that of Stereogenys and
different from that of Podocnemis is indisputable, but Swinton's
remark that the dorsal and lateral aspects of Shweboemys are
virtually identical to the comparable regions of Podocnemis and
unlike those of Stereogenys is an overstatement. Swinton (1939:
550) evidently compared the type of Shweboemys pilgrimi only
with Andrews' original description (1901: 442) of the type skull
of Stereogenys cromeri, which is somewhat crushed dorso-ventrally
in the facial region as Andrews himself noted (1901 : 443n). An-
other specimen (BMNH — R.3189), which Andrews later (1906:
301 and plate 25, fig. 1 ) referred to this species, is not flattened'
and reveals that virtually all of the characters used by Swinton
(1939: 550) to differentiate Stereogenys from Shweboemys — the
relative positions of the orbits and external nares, shape of the
orbits, and breadth as well as flatness of the skull — are artifacts
of preservation rather than taxonomically significant features.
Swinton also claimed (1939: 550) that the facial bones of
Stereogenys were more slender than those of Shweboemys. The
difference in thickness is easily explained, however. The type of
Shweboemys pilgrimi is considerably larger than that of Stereogenys
cromeri (12.5 versus slightly more than 8 centimeters from the
snout to the occipital condyle) and the thicker bones merely reflect
greater size. In dorsal and lateral aspects, therefore, the skuU of a
Shweboemys does not resemble that of Podocnemis more than it
does that of Stereogenys.
1 If anything, in terms of distortion, this skull may perhaps be some-
what compressed laterally.
1970 ASIATIC FOSSIL PELOMEDUSIDAE 9
Shweboemys pilgrimi Swinton 1939
(Plates I, IIA, IIIA, IVA)
Type. GSI 17255, an incomplete skull. The original description
of this specimen was accompanied only by some rather crude
sketches (Swinton, 1939, text-figures 1 and 2). Photographs of
the type showing comparable views are therefore reproduced as
Plate I.
Hypodigm. The type and BMNH— R. 8432, a slightly more
complete skull lacking the anterior portions of the premaxillae,
both ectopterygoid processes, part of the left and all of the right
paroccipital process, and, to whatever extent it may have been
developed, the supratemporal roof.
Horizon and locality. Pliocene or Pleistocene, Irrawaddy beds,
one mile NNE of Mauktet, Shwebo District, Burma.
Swinton's determination of a Pliocene age for the type (1939:
548) is questionable because he failed to state — perhaps because
the information did not exist — at what level within the Irrawaddy
sediments the specimen was found. It is generally recognized that
there are two faunal horizons within the Irrawaddy beds, a lower
one of Pliocene age and an upper one of Pleistocene age (Stamp,
1922: 498; Colbert, 1938: 267; Krishnan, 1960: 554). Un-
determined chelonian remains have been reported from the lower
beds near Yenangyaung (Stamp, 1922: 498), but these have never
been formally described. Fossil turtle fragments have also been
recovered from the upper Irrawaddy beds (Colbert, 1943: 417).
One of these, MCZ 1890 (MCZ 6305 in Colbert) represents the
left epiplastron of a very large tortoise while another (ANSP
14644)^ according to Colbert, may be a trionychid. No pelome-
dusid remains have been recognized among these specimens.
Whether the Irrawaddy beds of the Shwebo District, whose admin-
istrative center, the municipality of Shwebo, lies some 130-140
miles to the northeast of Yenangyaung, represent only the upper
part or the lower part of this sedimentary unit, or a mixture
of both, is unknown. Consequently, the absence of any reasonably
precise stratigraphic data for Swinton's specimen does not permit
a decision as to its age. Unfortunately, nothing is known about
the provenance, other than "Burma," of BMNH — R. 8432. Its
morphological identity with the type of 5. pilgrimi leads me to
believe that the two skulls are of essentially the same age.
10 BREVIORA No. 357
Emended diagnosis. Interorbital width slightly greater than di-
ameter of orbits; orbits circular, directed forward; maxillary tomia
curving upwards toward midline to form broad, semicircular notch;
medial borders of palatine flanges nearly parallel to each other as
far back as the opening for internal nares; little or no contact be-
tween pterygoids at midline; precondylar fossa lunate; trigeminal
foramen facing antero-laterally, situated low on wall of brain case;
breadth between postero-lateral corners of palatines equal to 40
per cent of skull length from snout to occipital condyle.
Not only does the British Museum specimen provide new infor-
mation about parts of the skull that were not preserved in the
type, but it also permits an important correction of Swinton's
description. The snout region of BMNH — R. 8432 is little dam-
aged and it is possible to determine unequivocally that, contrary
to his belief, nasals were lacking, as in all pelomedusids.
The type as preserved is 4V2 inches (11.4 centimeters) long
according to Swinton. A comparable portion of the British
Museum specimen measures 9.3 centimeters in length. The total
length of this skull (from snout to occipital condyle) is approxi-
mately 10.2 centimeters. With this information it is possible to
calculate the total estimated length of the type skull, assuming that
there were no significant ontogenetic changes in proportion, as
12.5 centimeters. Such large skulls indicate that adult representa-
tives of Shweboemys pilgrimi must have been imposing creatures,
roughly equivalent in size to Podocnemis expansa, the largest of
the living pelomedusid species.
The most persuasive indication that Swinton adduced to support
his contention that Shweboemys was a pelomedusid is its palatal
structure, which resembles that of Stereogenys more than that of
any other known turtle. This, however, is not a diagnostic
character, since it occurs elsewhere within the family only in
Bothremys and is rather widespread among cryptodires.
For taxonomic purposes, therefore, the most significant addi-
tional information provided by the British Museum skull is the
evidence that enlarged carotid channels were present (Fig. lA).
Such structures are known only in the pelomedusid genera
Podocnemis and Stereogenys and are not known in any other
turtle group. Much more substantial grounds now exist, con-
sequently, to confirm Swinton's belief that Shweboemys is truly a
pelomedusid.
1970
ASIATIC FOSSIL PELOMEDUSIDAE
11
Figure 1. Palatal views of: A — Shweboemys pilgrimi (BMNH — R.
8432); B— Shweboemys gaffneyi (BMNH— R. 8570). Solid parallel lines
represent areas of breakage. Matrix is indicated by randomly arranged
Vs. Stippled areas cover region in which thin sheets of surface bone have
broken off, thus making exact determination of position of sutures difficult.
Abbreviations: PM = premaxilla; M = maxilla; Pal = palatine; Pt =
pterygoid; Bs = basisphenoid; Bo = basioccipital; Eo = exoccipital; So =
supraoccipital; Etp = ectopterygoid process; cc = carotid channel.
12 BREVIORA No. 357
There does not appear to be any contact between the pterygoids
at the midHne, although less than perfect preservation on the
ventral surface of the basicranium of BMNH — R. 8432 necessi-
tates consideration of the possibility that these bones may have
barely met in an undamaged specimen (see Plate II A and Fig.
lA). Even if the pterygoids actually did meet at the midline,
their junction was clearly not extensive.^ Among pelomedusids,
this particular configuration is found only in this species and its
African relative, Shweboetuys antiqua. Another distinctive char-
acter is the position of the foramen for the trigeminal nerve
(Plate IIIA), which Swinton (1939: 551) was unable to detect
in the type specimen because of poor preservation. Instead of
being situated above the floor of the brain case and directed later-
ally, as in nearly all other pelomedusids, it is positioned much
lower and faces antero-laterally. In Bothremys cooki this foramen
is evidently situated as far down on the side of the brain case as
in Shweboemys pHgrimi, but it does not seem to have been directed
antero-laterally (Galfney and Zangerl, 1968: 220, figs. 13, 14,
and 16).
Breakage of the anterior ends of the premaxillae prevents an
exact determination of the shape of the external nares (Fig. 2A).
As preserved, the narial opening is ellipical, with its transverse
axis the longest. Damage to this same region also leaves some
question as to the actual shape of the upper jaw. Conceivably, it
may have had a downward curving beak, as in adult specimens of
Pelusios niger, or have simply been notched, as in many other
pelomedusids. Alternatively, its present rounded contour may
actually reflect its original shape. Whatever the case, it is clear
that there was a strong median indentation of some kind at the
midline.
Although the scroll-like outer portion of the laterally projecting
ectopterygoid processes have not been preserved in either speci-
men of Shweboemys pilgrimi, these structures did exist. Evidence
to this effect is preserved on BMNH — R. 8432, where the basal
portion of these protuberances can be seen on both sides.
1 Contrary to Swinton's statement (1939: 550), no portion of the
pterygoids is preserved on the holotype. What he evidently interpreted as
the palatine-pterygoid suture appears to be a transverse crack across the
ventral surface of the palatines (Plate lA).
1970
ASIATIC FOSSIL PELOMEDUSIDAE
13
3 CM
3 CM
Figure 2. Facial views of: A — Shweboemys pilgrimi (BMNH — R.
8432); B—Shweboemys gaffneyi (BMNH— R. 8570). Solid parallel lines
represent areas of breakage.
♦
14
BREVIORA
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1970 ASIATIC FOSSIL PELOMEDUSIDAE 15
Scute sulci can be detected on the roof of the British Museum
skull (Fig. 3). The anterior border of the interparietal overlapped
slightly onto the postero-lateral corners of the frontal bones and
was bow-shaped, more like that of Podocnemis sextiiberculata
{cj. Siebenrock, 1902, fig. 9) than any other pelomedusid known
to me. Behind the orbit a scute intervened between the maxillary
and frontal scutes. Whether this was an enlarged masseteric scute,
as in Podocnemis diimeriliana, or a subocular scute, as in most of
the other species of Podocnemis (Williams, 1954), is uncertain.
Shweboemys gaffneyi sp. nov.^
(Plates MB, IIIB, IVB)
Type. BMNH — R. 8570, a partial skull lacking the premaxillae,
most of the prefrontals, the bones of the cheek region, the quad-
rates and squamosals, the supraoccipital crest, and whatever
supratemporal roofing there may have been.
Hypodigm. The type, only.
Horizon and locality. Early Miocene, Bugti Hills, Baluchistan,
West Pakistan.
No locahty or stratigraphic data are associated with this speci-
men, but apparently it was obtained by C. Forster-Cooper during
one of his two expeditions to the area around Dera Bugti during
the years 1910 and 1911 and would therefore be of the age and
from the general region cited above. It was subsequently given to
Professor D.M.S. Watson for description, who in turn passed it
on to Dr. E. E. Williams for the same purpose. Other commit-
ments having prevented either from formally describing this
interesting skull, the task has now fallen to me.
Pilgrim (1908: 144) briefly mentioned that chelonian remains
are not uncommon in the Miocene sediments of the Bugti region.
Unfortunately, none of these have ever been described. Knowl-
edge of their existence nevertheless reinforces the probabihty that
the skull under consideration is of the age of and from the locality
given above.
Diagnosis. Differing from S. pilgrimi in : interorbital width less
than diameter of orbits; tomial margins of upper jaw (so far as
preserved) horizontal, not notched; medial borders of palatine
1 The species is named for Dr. Eugene Gaffney in recognition of his
work on pelomedusid turtles.
16 BREVIORA No. 357
flanges curving away from midline; breadth between postero-
lateral corners of palatines equal to 50 per cent of skull length
from snout to occipital condyle; broad contact between pterygoids
at midline; precondylar fossa semicircular; foramen for trigeminal
nerve situated relatively higher on ascending wall of brain case
and directed laterally. Differing from S. antiqua in: lack of fore-
head groove; orbits round rather than oval; no median notch in
upper jaw; broad contact between pterygoids at midline; size
much greater.
In total length, this skull is slightly longer than the smaller of
the two specimens of Shweboemys pilgrimi (10.5 versus 10.2
centimeters for the distance from the snout to the occipital
condyle). The width at the postero-lateral corners of the palatines
is significantly greater, however, indicating that the skull of S.
gaffneyi was proportionately wider. Compared to S. pilgrimi, the
orbits of the new species are also proportionately much larger.
The increase has been achieved primarily by dorsal emargination
of the external face of the maxilla. The diameter of the orbits in
S. pilgrimi is essentially the same as the distance across the maxilla
from the base of the orbit to the tomium. In contrast, the diameter
cf the orbits in S. gaffneyi is nearly twice as great. Because the
premaxillae are missing, it is impossible to reconstruct the shape
of the upper jaw at the midline. There may have been some kind
of median notch, but if so it must have been relatively small and
thus quite unlike that of S. pilgrimi (see Fig. 2B).
The two Asiatic species differ somewhat in the structure of their
secondary palates. In S. gaffneyi, the secondary palate is broader
than it is long, whereas in S. pilgrimi this region is slightly longer
than wide (cf. Figs. lA and B). For analyzing proportional dif-
ferences, direct comparisons may be made between comparable
measurements of the two British Museum skulls of Shweboemys,
each representing one of the Asiatic species, since they are both
of nearly the same length and are equally well preserved in the
region under consideration. At the point where the sutures be-
tween the maxillae and the palatines reach the outermost extent
of the secondary palates, the distance across the secondary palate
of the type of S. gaffneyi is 7.2 centimeters, whereas this same
distance in BMNH — R. 8432 is 6.1 centimeters. But the length
of this structure is 5.3 centimeters in the former and 6.7 in the
latter. Another difference, of unknown biological significance,
involves the pitting on the ventral surface of the secondary palate.
1970 ASIATIC FOSSIL PELOMEDUSIDAE 17
In both species, these pits appear to be more abundant on the
maxillae than on the palatines. The maxillary pits, however, seem
to be deeper, larger in diameter, and fewer in number in 5.
gaffneyi (see Plate II).
No scute sulci are clearly discernible on the preserved part of
the skull roof of 5. gaffneyi. Linear depressions, which might be
interpreted as scute furrows, follow the courses of the fronto-
parietal and fronto-postorbital bone sutures. But a similar groove
running antero-posteriorly along the midline does not have a
homologue in any other pelomedusid. Thus I am dubious that any
of these indentations necessarily corresponds in position to the
actual boundaries between scutes.
The position of the trigeminal nerve foramina and the extent
to which the pterygoids meet on the ventral surface of the skull in
5. gaffneyi are typically pelomedusid and clearly serve to differenti-
ate this species from 5. pilgrimi, which is specialized in these
characters. The skull of S. gaffneyi exhibits no features that would
bar the species from the ancestry of S. pilgrimi.
The skull of S. gaffneyi differs from that of 5. antiqua in a num-
ber of respects, of which a few may be mentioned here, pending
detailed description of the Fayum species. Size is the most obvious
(although not necessarily the most significant taxonomically ) dif-
ference between the two: the distance from the snout to the
occipital condyle in S. gaffneyi is more than one and one-half
times longer than in 5. antiqua ( 10.5 versus 6.6 centimeters). The
orbits of S. gaffneyi are round and directed forward, those of S.
antiqua are oval and face laterally. There is a forehead groove
between the orbits of the latter but not of the former. As in S.
pilgrimi, and in strong contrast to 5. gaffneyi, there is little or no
contact between the pterygoids at the midline in 5. antiqua.
DISCUSSION
Of the small number of Asiatic fossil pelomedusids hitherto
described, one, "Podocnemis" indica, is not well enough known
at the present time to merit formal taxonomic assignment. How-
ever, Carteremys probably was a pelomedusid, and Shwebuemys
certainly was.
The relationships of Carteremys within the Pelomedusidae are
uncertain. On the basis of skull structure, Shweboemys appears
to have been more closely related to Stereogenys than to any other
18 BREVIORA No. 357
pelomedusid. Within the genus, Shweboemys pilgrimi may well
have been derived more or less directly from 5. gaffneyi, and
there is no reason why this latter species could not in turn have
been directly descended from S. antiqua. Occurrences of this
genus are separated by such great distances and represented by
such a paucity of material, however, that future discoveries may
reveal that this interpretation is overly simplistic.
Shweboemys is the only pelomedusid genus so far known to
have established a successful, enduring lineage outside of Africa
or South America. The Shweboemys lineage appears to have been
restricted to southern Asia and Africa; no pelomedusid has thus
far been reported, even questionably, from central or eastern Asia.
The localities where the two Asiatic species of this genus occur
are very widely separated geographically and no representative
has been recorded from the intervening (and considerably
younger) Siwalik deposits of India. Nothing, however, has been
published on Siwalik fossil turtles for more than three-quarters of
a century. Those described by Lydekker (1885), apart from the
large testudinids, show relationships to the recent turtle fauna of
India, but available collections need to be examined thoroughly
with an eye to the possible presence of pelomedusid remains.
Some inferences are possible concerning the ecology of the two
Asiatic species of Shweboemys: the Irrawaddy beds are fluviatile
in origin (Krishnan, 1960: 498), so that this species was evidently
not a marine form.^ In view of the fact that all pelomedusids
(except for one or possibly two undescribed fossil genera from
Africa) are aquatic, S. pilgrimi was probably a freshwater rather
than a terrestrial turtle. Pilgrim (1908: 159) referred to the
Bugti beds from which S. gaffneyi was presumably recovered as
a '^freshwater formation" and Krishnan (1960: 492) regarded
them as being fluviatile, so that this species was in all likelihood
also a freshwater rather than a marine form. Moreover, the
specialized palatal structures of these two species have definite im-
plications regarding their feeding habits. Like some living triony-
chids with enlarged secondary palates, their diet may have con-
sisted largely or perhaps even exclusively of molluscs of one sort
or another.
1 1 am suggesting elsewhere that the pelomedusids were of marine origin.
1970 ASIATIC FOSSIL PELOMEDUSIDAE 19
ACKNOWLEDGMENTS
I am particularly grateful to Mr. C. A. Walker of the British
Museum (Natural History) for bringing to my attention the skull
of Shweboemys pilgrimi belonging to that institution. To the
authorities of the British Museum (Natural History) I am indebted
for permission to describe this specimen as well as the type of
S. gaffneyi. I am also much obliged to Professor Bryan Patterson
and Dr. E. E. Williams for critically reading this manuscript, to
Professor B. Kummel for help concerning the stratigraphy of West
Pakistan and Burma, and to Mr. A. D. Lewis for his skillful
preparation work on both of the British Museum skulls. The
photographs of the type of 5. pilgrimi reproduced in Plate 1 were
sent to Dr. E. E. Williams by the authorities of the Geological
Survey of India. The figures were drawn by Mr. Laszlo Meszoly.
Finally, I would like to express my appreciation to the National
Geographic Society for their generous financial support of my
research on pelomedusid turtles.
LITERATURE CITED
Andrews, C. W. 1901. Preliminary note on some recently discovered
extinct vertebrates from Egypt. (Part II.) Geol. Mag., 8: 436-444.
. 1906. A descriptive catalog of the Tertiary Vertebrata of
the Fayum, Egypt. Brit. Mus. (Nat. Hist.), London. 324 pp.
Carter, H. J. 1852. Geology of the Island of Bombay. Jour. Bombay
Branch Roy. Asiatic Soc, 4, 16: 161-215.
Colbert, E. H. 1938. Fossil mammals from Burma in the American
Museum of Natural History. Bull. American Mus. Nat. Hist. 74, art.
6: 255-436.
1943. Research on early man in Burma, Part III:
Pleistocene vertebrates collected in Burma by the American Southeast
Asiatic expedition. Trans. American Phil. Soc, n.s., 32, part 3:
395-429.
Gaffney, E. S.. and R. Zangerl. 1968. A revision of the chelonian
genus Bothremys (Pleurodira: Pelomedusidae), Fieldiana (Geol.), 16
(7): 193-239.
Krishnan, M. S. 1960. Geology of India and Burma (4th ed.). Madras.
Higgenbothams (Private) Ltd. 604 pp.
Lydekker, R. 1885. Indian Tertiary and post-Tertiary Vertebrata.
Siwalik and Narbada Chelonia. Mem. Geol. Surv. India, Palaeont.
Indica, sen 10, 3, part 6: 155-208, pis. 18-27.
1887. Indian Tertiary and post-Tertiary Vertebrata.
Eocene Chelonia from the Salt Range. Mem. Geol. Surv. India,
Palaeont. Indica, ser. 10, 4, part 3: 59-65, pis. 12-13.
20 BREVIORA No. 357
1890. Note on certain vertebrate remains from the Nagpur
District. Records Geol. Surv. India, 23: 20-24.
Pascoe, E. H. 1964. A manual of the geology of India and Burma,
3 (3rd ed.). Geol. Surv. India: XXIV and pp. 1345-2130.
Pilgrim, G. E. 1908. The Tertiary and post-Tertiary freshwater de-
posits of Baluchistan and Sind, with notices of new vertebrates.
Records Geol. Surv. India, 37, part 2: 139-166, pis. 2-4.
Robinson, P. L. 1970. The Indian Gondwana formations — a review.
l.U.G.S. First Symposium on Gondwana Stratigraphy: 201-268.
RoMER, A. S. 1956. Osteology of the Reptiles. Chicago, The University
of Chicago Press. 772 pp.
SiEBENROCK, F. 1902. Zur systematik der schildkroten-gattung Podoc-
nemis Wagl. Sitz. der kaiserl. Akad. der Wiss. in Wien, Math.-Naturw.
Classe, 111, abt. 1: 1-14.
Stamp, L. D. 1922. An outline of the Tertiary geology of Burma. Geol.
Mag., 59 (11): 481-501.
SwiNTON, W. E. 1939. A new fossil fresh-water tortoise from Burma.
Rec. Geol. Surv. India, 74, pt. 4: 548-551.
Wadia, D. N. 1953. Geology of India (3rd ed.). London, Macmillan
and Co. 531 pp.
Williams, E. E. 1953. Fossils and the distribution of chelyid turtles,
1. "Hydraspis leithii" (Carter) in the Eocene of India is a pelomedusid.
Breviora, no. 13: 1-8, pis. 1-3.
^__ 1954. A key and description of the living species of the
genus Podocnemis {sensii Boulenger) (Testudines, Pelomedusidae).
Bull. Mus. Comp. Zool., Ill, (8): 279-295.
Zangerl, R. 1969. The turtle shell. In Biology of the Reptilia, /.
Morphology A (C. Gans, ed.) London and New York, Academic Press.
311-339.
970
ASIATIC FOSSIL PELOMEDUSIDAE
21
-'-r^np^
.^i*^^*^"^'
»>•
B
I. Type specimen oi SJnveboemys pilgrimi (GSI 17255): A —
palatal view of skull; B — dorsal view. Approximately Va natural
size.
22
BREVIORA
No. 357
!
■
1 11
Hj CM
■
■1
^^^^1 • •'\^' "
A
*^"^^^^§L^^-
^'' -«<■•■ >^^o|^^3H|^t^
k
b
"** "*** ^^BSiBt
ilraMI^T'^
^^^^s
»^«w
■J^
■J^^^Pk
'^^^^^^m^B^^-, «-v» --t^^t "•» '^^H
'
II. Palatal views of: A — Shweboemys pilgrimi (BMNH-
R. 8432); "R— Shweboemys gafjneyi (BMNH— R. 8570).
1970
ASIATIC FOSSIL PELOMEDUSIDAE
23
III. Lateral views of: A — Shweboemys pilgrimi (BMNH-
R. 8432); B— Shweboemys gaffneyi (BMNH— R. 8570). tgf
trieeminal nerve foramen.
24
BREVIORA
No. 357
CM
IV. Dorsal views of: A — Siiweboemys pilgiimi (BMNH — R.
8432); B—Shweboemys gaffneyi (BMNH— R. 8570).
BREVIORA
Miaseium of Comparative Zoology
Cambridge, Mass. 30 November, 1970 Number 358
South American anoles: Anolis apollinoris Boulenger 1919,
a relative of A. biporcatus Wiegmann (Sauria, Iguanidae)
Ernest E. Williams
Abstract. Anolis apollinaris is a central Andean derivative of A. bipor-
catus, probably from an earlier invasion of South America than that which
has provided the present Colombian, Ecuadorian, and western Venezuelan
populations of the latter species.
Anolis apollinaris Boulenger 1919 was described from a unique
type, a female, said to come from "near Bogota." The description
made no mention of relationships.
The next mention of the species was made by Burt and Burt
(1931: 255), who referred numerous Colombian specimens in the
American Museum to this species. They suggested that the species
belonged to the "chrysolepis stock" but also said that their speci-
mens closely resembled A. gemmosus of Ecuador with which they
believed A . apollinaris "may prove to be identical or subspecifically
allied." An examination of the type of apollinaris in the British
Museum shows that these statements of relationships are entirely
mistaken and that the specimens referred to the species by Burt
and Burt — one specimen received in exchange from the American
Museum by the Museum of Comparative Zoology — are mis-
identified.^
1 The type of A. gemmosus O'Shaughnessy has also been examined. It
is not of chrysolepis stock nor related at all closely to the two species mis-
identified by Burt and Burt as A. apollinaris. The affinities of A. gemmosus
are with A. fasciatus Boulenger and A. andianus Boulenger.
2 BREVIORA No. 358
A correct judgment on the affinities of A. apollinaris was made
by E. R. Dunn in 1944 (p. 25), who at that time reported:
"The Instituto de La Salle has a specimen of this lizard (de-
scribed from 'near Bogota') from Paime, Cundinamarca, 1038
meters. A number of students have overlooked the statement that
this is a large Anolis (type head-body length 106 mm) and mis-
applied the name. Thus the "Anolis apollinaris" of Burt and Burt
(1921 I sic J, p. 255) is not Boulenger's species but a composite of
two smaller species, incomperliis Barbour from Villavicencio and
mariaruin Barbour from Medellin^ True apollinaris is allied to
solijer of Santa Marta and copei of Central America."
The two latter names are now regarded as synonyms of bipor-
catus (see Williams, 1966) and it is with this species, which ranges
from Mexico to Ecuador, that apollinaris requires comparison.
Brother Niceforo Maria of the Instituto La Salle tells me that
Dunn's specimen of A. apollinaris was one of many specimens
destroyed in a fire at the Institute in 1948. Fortunately, a number
of previously unreported specimens have been discovered, one in
the Institut Royale (Brussels), a series in the Zoologische Staat-
sammlung (Munich) and three, indeed, in more recent collections
of the Instituto La Salle (ILS), and two more in the American
Museum of Natural History (AMNH).
On the basis of these new specimens and the type specimen at
the British Museum (BM), I present a revised standard descrip-
tion of the species:
Anolis apollinaris Boulenger
Type. BMNH 1919.3.6.7 (1946.8-13.22), from near Bogota,
Cundinamarca, Colombia.
Referred specimens. (All Colombia.) Antioquia (all Cauca
Valley): AMNH 38725, Sabanalarga; ILS 81, Puerto Antioquia.
Caldas: ILS 101, Pueblo Rico. Cundinamarca: Brussels 3580, La
Esperanza, 1250 m; ILS 65, Paime; ILS 106, Quipile; Munich
427-432, San Pablo, west side of cordillera between Bogota and
La Dorada. "Western Colombia": AMNH 4844.
1 A. incompertus Barbour is a composite species: specimens from
Villavicencio are A. chrysolepis scypheus Cope and, from near Bogota. A.
tropidogaster Hallowell. A. mariaruin Barbour is a synonym of A. antonii
Boulenger. All types have been examined.
1970
ANOLIS APOLLINARIS
u
d
Z,
B
Xi
o
c
3
2
BREVIORA
No. 358
Diagnosis. Allied to biporcatus Wiegmann and its subspecies
parvauritus Williams but differing in color, in one or no scales
separating nasal from rostral, and in a modally higher number of
lamellae under phalanges ii and iii of the fourth toe.
Head. Head scales small, sharply uni- or tricarinate. Ten to
thirteen scales across snout between second canthals. A distinct
frontal depression, scales within it not smaller than surrounding
scales. Five to nine scales border rostral posteriorly. Circumnasal
scale separated from rostral by one small scale or in contact. Six
to seven scales between circumnasals dorsally.
Supraorbital semicircles separated from each other by 2-4 scales,
from the supraocular disk by one row of smaller scales. Supra-
ocular disk not very distinct, of 4-12 keeled scales grading laterally
into granules. One to three overlapping elongate supraciliary scales,
continued posteriorly by granules. Anterior corner of supraocular
filled by larger subgranular scales. Canthus sharp, of 6-7 over-
lapping scales, the first and second or second and third the largest.
Five to seven loreal rows, subequal or the uppermost largest. Tem-
poral scales granular. A distinct double line of enlarged inter-
temporal scales. Supratemporals granular, slightly smaller than
Figure 2. Anolis apollinaris Munich No. 422. Dorsal view of head.
1970 ANOLIS APOLLINARIS 5
temporals. Scales surrounding interparietal moderately to abruptly
enlarged, swollen, largest anteriorly and laterally. Interparietal less
than or greater than ear, separated from semicircles by 3-4 scales
on each side.
Suboculars separated from supralabials by one row of scales (or
narrowly in contact), anteriorly separated from canthal ridge by
one scale, posteriorly continued by an indistinct double row of
smaller scales. Seven to eight supralabials to center of eye.
Mental slightly wider than long, in contact with 4-8 scales be-
tween supralabials posteriorly. Sublabials not well differentiated.
Central throat scales quadrangular, swollen, gradually increasing
in size laterally.
Dewlap. Dewlap in male large with close-packed scales. A
gular fold only in female, moderate, scales rather closely packed.
Trunk. Middorsals slightly enlarged, swollen, keeled. Dorsal
and flank scales keeled, subequal. Ventrals larger, weakly keeled,
imbricate, not mucronate.
Limbs. Largest fore and hind limb scales strongly unicarinate,
except at knee and elbow, smaller than largest ventrals. Supra-
digital scales multicarinate. Twenty-four to twenty-seven lamellae
under phalanges ii and iii of fourth toe.
Tail. Slightly compressed, almost evenly scaled all round. Ver-
ticils indistinct. All scales keeled. Enlarged postanals in male.
Size. Type: 106 mm snout-vent length.
Comparison. Table 1 lists the major features differentiating A.
apollinaris and A. biporcatus. I comment on each of these features
below:
1. Scales in narial area. The exact pattern of the scales sur-
rounding the naris and their relation to the rostral have been
repeatedly used in lizards generally (e.g., geckos), and this pattern
has also proved empirically very useful at the species level in
Anclis. Published examples of the utility of this character at the
species level are Ruibal and Williams (1961 ) and Lazell (1964).
Although, like all squamation characters in Anolis, these patterns
are subject to some intraspecies variation, they are rather surpris-
ingly constant. Frequently a single pattern is consistently main-
tained; this is usually a simple one. More complex patterns tend
to greater variation but the variations are readily derivable from
the modal condition (Fig. 3). The pattern of apollinaris is always
simpler than that of biporcatus and may be more primitive. (The
judgment that this pattern may be more primitive is based not upon
BREVIORA
No. 358
Figure 3. Nasal rostral relationships. Upper left: Anolis biporcatus
biporcatiis, MCZ 15426. Upper right: A. b. parvaiiritus, MCZ 78942.
Lower left: A. apollinaris, Munich 422. Lower right: A. fraseri, MCZ
43772.
its simplicity but upon its association with other characters re-
garded as primitive and on its occurrence in species regarded on
other grounds as primitive.) The variation in apollinaris is in the
direction of the pattern in biporcatus, but there is no overlap. This
is a sharp and clear distinction.
2. Supraciliary scales (Fig. 4). Again the pattern of this area
tends to be species specific and again the condition in apollinaris
tends to be simpler and perhaps more primitive than that of bipor-
catus. The common pattern in biporcatus is indeed unusual (though
not unique). Two patterns are common for the supracihary region
of anoles: (1) one or more elongate supraciliaries followed by un-
differentiated granules; (2) one or more elongate supraciliaries
followed by a double series of enlarged scales rather even in size.
970
ANOLIS APOLLINARIS
Figure 4. Supraciliary area. Top: Anolis apoUinaris, Munich 422.
Lower left: A. b. biporcatiis, MCZ 15426. Lower right: A. fraseri, MCZ
43772.
A. biporcatiis, exhibiting several rather short supraciharies grading
into large scales that tend to grade again into granules, presents a
condition hardly more frequent than that of A . fraseri with its series
of quadrate scales along the whole supraciliary margin, the first of
these sometimes elongate.
3. Scales around interparietal. In general, the degree of
enlargement of scales around the interparietal is a good specific
character in Anolis. Particularly valuable may be the degree of
enlargement of the scales posterior to the interparietal as compared
with the adjacent dorsal or supratemporal scales. There may be
rapid intergradation of enlarged scales lateral to the interparietal
into much smaller dorsal and supratemporal scales, or the scales
behind the interparietal may be sharply and conspicuously larger
than dorsal or supratemporal scales (as in some apoUinaris). The
two subspecies of A. biporcatus difl'er in this regard. A. apoUinaris
is variable; perhaps the variation is geographic, but there is not
enough material to say.
4. Ear shape and position. The ear of apoUinaris is quite dif-
ferent from that of either southern or northern biporcatus. It is
8 BREVIORA No. 358
closer in size to that of southern biporcatus but quite distinct in its
obliquity, a rather unusual feature.
5. Ventral keeling. This character is minor. Keeled ventrals
may be a good species character, but there are many instances of
intraspecies variability, both geographic and (typically qualitatively
less extreme) at a single locality. A. apollinaris has the ventrals
more weakly keeled than either subspecies of biporcatus.
6. Toe lamellae. The number of toe lamellae is an extremely
useful character in Anolis and very characteristic of species. It is,
however, subject to variability (a range of 6 or 7 is quite usual)
and overlap is, as in the present case, frequent. A. apollinaris
tends to a higher number of toe lamellae than either subspecies
of biporcatus.
7. Color. I have no descriptions of color in life of apollinaris
and the varying colors of biporcatus as preserved (it is uniform
green in life) do not make comparison very easy. Boulenger de-
scribed the type female as "Dark olive above and on the sides,
with a fine blackish network, head and a vertebral band pale, the
latter with narrow transverse processes; small round light spots on
the sides and tail; forearm, tibia and lower parts pale green."
The Brussels specimen has preserved its pattern rather well.
Description follows: Head greyish. A dark streak from back of
eye to shoulder, there merging with dark flanks. Below this, labials
and nape lighter, their color continuous with the smudged grey of
the throat. A hght brown middorsal zone, irregularly darker lat-
erally, bordered on each side by a narrow grey line. Flanks dark
brown with indications of white spots or broken narrow vertical
white bars. Forelimbs obscurely annulate, hind limbs boldly so.
Tail above with longitudinally oval light spots with irregularly dark
centers. Belly lighter than any part of dorsum but still heavily
infuscated. Tail below lighter still.
The new specimens resemble the Brussels specimen in head
coloration, as the British Museum type now does; presumably the
latter differs from Boulenger's description as a result of change
during preservation.
A. apollinaris, when compared with A. biporcatus, differs in few
and superficially trivial ways. The ventrals are less strongly keeled.
There are fewer scales between nostril and rostral: one or more
rather than two or three. The color is quite unlike anything 1 have
seen in biporcatus. I am neither confident that this form is a full
1970 ANOLIS APOLLINARIS 9
species nor convinced that it is not. It is certainly not to be con-
fused with Mexican or Central American biporcatus, nor with the
south Colombian-Ecuadorian population, nor does it resemble the
two Venezuelan specimens of that species. Whether it is distinct
from, or intergrades with, some of the other Colombian or the
Ecuadorian populations is a matter for future discovery. Provi-
sionally, since it was described as a species, it may remain so
allocated.
A. biporcatus has the distribution of a recent immigrant into
South America, extending from Panama through the Choco region
of Colombia down into Ecuador west of the Andes and extending
eastward through the Santa Marta Mountains into western Vene-
zuela. A. apollinaris, now recorded from Antioquia, Caldas, and
Cundinamarca, may be supposed to have arisen from A. bipor-
catus by isolation and subsequent minor modification in the central
Andean regions of Colombia. Since it has some primitive features
(e.g., nasal-rostral relationship) and since it has reached species
status, it may represent an earlier invasion of South America than
that which resulted m A. b. parvauritus Williams and the Vene-
zuelan specimens oi A. b. biporcatus.
REFERENCES
BouLENGER, G. A. 1919. Descriptions of two new lizards and a new frog
from the Andes of Colombia. Proc. Zool. Soc. London, 1919: 79-80.
Burt, C. E., and M. D. Burt. 1931. South American lizards in the col-
lection of the American Museum of Natural History. Bull. American
Mus. Nat. Hist. 61: 227-395.
Dunn, E. R. 1944. Herpetology of the Bogota area. Revista Acad. Co-
lombiana Cienc. 6: 68-81.
Lazell. J. D., Jr. 1964. The anoles (Sauria, Iguanidae) of the Guade-
loupeen Archipelago. Bull. Mus. Comp. Zool. 131: 359-401.
RuiBAL. R., AND E. E. Williams. 1961. The taxonomy of the Anolis
honwlechis complex of Cuba. Bull. Mus. Comp. Zool. 125: 209-246.
Williams, E. E. 1966. South American anoles: Anolis biporcatus and
Anolis fraseri (Sauria, Iguanidae) compared. Mus. Comp. Zool.,
Breviora No. 239: 1-14.
(Received 9 June 1970.)
10
BREVIORA
No. 358
■10
O'
-:^/0'
Figure 5. Map of the distribution of Anolis apollinaris in central
Colombia.
1970
ANOLIS APOLLINARIS
11
TABLE 1
apollinaris b. biporcatus
scales across snout 9-13 7-11
b. parvauritus
8-13
scales between nasal
and rostral
0-1
2-3
2-3
scales between supra-
orbital semicircles
2-4
1-4
0-3
supraciliaries
1-2 elongate plus usually 3-4 short as in biporcatus
series of small supraciliaries
scales of rather plus a series of
uniform size small scales of
variable size
scales behind
interparietal
variable, slightly abruptly larger
to abruptly larger than dorsals
than dorsals
grading gradually
into dorsals
scales separating
interparietal from
semicircles
3-5
3-6
3-7
loreal rows
5-8
5-10
6-9
supralabials to
center of eye
ear
7-8
8-11
small to moderate moderate to
large, vertical
7-12
small
ventrals
weakly keeled
strongly keeled,
mucronate
strongly keeled,
mucronate
lamaellae under
phalanges ii and iii
of fourth toe
25-27
22-26
22-26
BREVIORA
MiaseiLaitii of Comparative Zoology
Cambridge, Mass. 30 November, 1970 Number 359
The Swimbladder as a Juvenile Organ
in Stromateoid Fishes
Michael H. Horn^
Abstract. The swimbladder regresses in 14 of the 15 genera of strom-
ateoid fishes and apparently in each case before maturity is reached. In
one genus the swimbladder is absent. The organ appears to be completely
functional and is present in juvenile fishes that inhabit the surface layers
of the ocean, often in association with jellyfish medusae or floating objects.
In the transition from the juvenile to the adult habitat and mode of life,
the swimbladder regresses to a nonfunctional state, and other morphological
changes occur.
A detailed study in progress of the state and structure of the
swimbladder in stromateoid fishes in relation to their evolution
and ecology shows that the organ regresses with age in 14 of the
15 genera comprising this suborder of perciform teleosts. The
regression of the swimbladder occurs at different ages, depending
upon the species in question and seems to be correlated with other
morphological changes and with certain changes in mode of life.
Thus, the state of the swimbladder may be useful in predicting
or determining at what age or size a particular species undergoes
a shift in habitat or way of life.
The suborder Stromateoidei consists of six families (Haedrich
& Horn, 1969), the members of which range in maximum size
from about 30 to about 120 centimeters. It is a fairly diverse
group of temperate and tropical marine fishes, which, as adults,
variously occupy a wide range of depths in coastal and oceanic
waters. The Stromateidae occur at all depths over the continental
shelf. The Ariommidae are benthic or benthopelagic on the shelf.
1 Present address: Department of Biological Science, California State
College, Fullerton, California 92631.
2 BREVIORA No. 359
The Centrolophidae are either coastal, as in Hyperoglyphe and
Seriolella, or oceanic at various depths, as in Centrolophiis and
Icichthys. The Nomeidae, Tetragonuridae, and Amarsipidae are
oceanic usually in epi- or mesopelagic layers. However, as occurs
in many other marine fishes, the larvae and juveniles of stroma-
teoids are pelagic in the surface layers, mostly in the upper 100
meters.
Stromateoid fishes commonly undergo marked changes when
approaching maturity, and these changes are often associated with
the migration from the surface layers to the deeper layers where
the fishes live as adults (Haedrich, 1969). Along with certain
changes in body proportions (see Haedrich, 1967; and Horn,
1970), including, frequently, changes in the length of paired fins,
the swimbladder regresses. This regression and the significance of
the swimbladder in the life of young fishes are discussed below.
MATERIALS AND METHODS
Specimens examined are from the collections of the British
Museum (Natural History); the Zoological Museum, Copen-
hagen; and, the Woods Hole Oceanographic Institution. The
fishes from the last institution will ultimately be deposited in the
Museum of Comparative Zoology, Harvard University. Specimen
data will be included in a forthcoming comprehensive paper on
stromateoid swimbladders.
Swimbladder volumes were calculated treating the bladder as
an ellipsoid. Body volumes were determined by displacement in
water. Ten per cent was allowed for shrinkage of the preserved
material, and volume determinations were made from swimblad-
ders that were in most cases well expanded. The number of retial
capillaries was estimated from thin sections of swimbladders.
RESULTS AND DISCUSSION
Very little mention has been made of the swimbladder in
stromateoids. Jordan & Evermann (1896) stated that the organ
was "usually absent" in the Stromateidae. Fowler (1936) indi-
cated that it was "present or absent" in his treatment of several
stromateoid genera. Grey (1955) reported it to be absent in
Tetragonurus. Haedrich (1967) cited its presence in Ariomma.
While I have not yet examined every species, I have found a
swimbladder to be present in all genera except Pampus.
1970
SWIMBLADDER OF JUVENILE STROMATEOIDS
The swimbladder in stromateoids (Fig. 1) is euphysoclystous,
i.e., with a distinct gas-resorbing area — a posterior chamber
separated from the anterior by a diaphragm — and has unipolar
retia that are either medial or posterior in position. It is generally
of a relatively simple type found in a number of perciform fishes
(N. B. Marshall, personal communication).
Figure 1. Ventral (slightly flattened) view of the swimbladder of
Nomeus gronovii, 26.4 mm SL; rm = rete mirabile; rv = retial vein;
ra = retial artery; gg = gas gland; ac = anterior chamber; dm = dia-
phragm; pc = posterior chamber. (X 33)
The swimbladder in these fishes is relatively small and fre-
quently below the 3.1-5.7 range of percentage volumes calculated
by Alexander (1966) to be necessary for neutral buoyancy in
sea water (Table 1). Measurements of swimbladder percentage
volume for Hyperoglyphe, Cubiceps, and Ariomma show the
organ to be within the above range, but in other genera, notably
Schedophiliis and Nomeus, the percentage volume may be as low
as 0.5 or 0.6 per cent. Some calculated values may be lower
than normal because of the swimbladder being in a slightly con-
tracted state. The organ does appear to be completely developed
and almost certainly functional. The blood supply is complete,
and the surface area of the gas gland and the length of the retia
in relation to swimbladder size (Table 1 ) compare closely with
the range of values calculated by Marshall (1960: tables 3, 4, &
5) for deep-sea fishes with well-developed swimbladders. The
organ is fully formed in the smallest individuals (9 mm SL) that
have been examined. Regression to a nonfunctional state occurs,
apparently in all species, before maturity is reached.
BREVIORA
No. 359
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1970 SWIMBLADDER OF JUVENILE STROMATEOIDS 5
Being fully developed and certainly capable of hydrostatic ad-
justment, the stromateoid swimbladder seems very unlikely to be
merely an evolutionary remnant. The clue to its significance
appears to lie in the behavior of the juveniles. The young of all
or nearly all of the species live in the surface layers and fre-
quently in association with jellyfish medusae or other animate or
inanimate floating objects. Protection is presumably provided by
the medusae or floating objects, and the fishes feed upon the
small invertebrates concentrated around the objects or upon the
jellyfishes themselves. This existence seems to require both con-
siderable maneuverability and the ability to hover and remain
motionless in midwater. I have observed young Peprilus triacan-
thus hovering beneath and near the tentacles of the sea nettle,
Chrysaora qiiinquecinha, and they show little locomotor activity
except for backing of water by the pectoral fins. Mansueti (1963)
has made similar observations on Peprilus alepidotus {= P.
paru.). Few, if any, of the stromateoid genera which consort
with medusae are completely immune to jellyfish toxins but merely
avoid the tentacles, according to observations made by Mansueti
(1963) on P. alepidotus, by Maul (1964) on Mupus {-Schedo-
philus), and myself on P. triacanthus. Lane (1960) reports that
Nomeus gronovii can survive doses of Physalia toxin as much as
ten times that which would kill other fishes of the same general
size and type; however, Nomeus still exhibits considerable agility
in avoiding the tentacles of Physalia.
The swimbladder, even if smaU, would provide a degree of
buoyancy, although not necessarily complete neutral buoyancy.
During the period of life in which the fishes have a gas bladder,
the skeleton is not well ossified and the musculature may not be
completely developed; thus, the juveniles probably have a lower
specific gravity than the adults. In those adults with soft muscula-
ture and light ossification, such as Schedophilus, the swimbladder
probably becomes unimportant and uneconomical because of a
change in mode of life, even though, as in juveniles, a small volume
of gas would provide nearly neutral buoyancy. Using Alexan-
der's (1966) formula for calculating the percentage swimbladder
volume necessary to achieve hydrostatic equilibrium, I find that
only a 1 per cent reduction in specific gravity of the fish lowers the
required percentage volume from 3.1 per cent, the lower figure
in Alexander's calculated range, to 2.2 per cent. The latter figure
is within or near the range of volumes for most of the stromateoid
6 BREVIORA No. 359
genera (Table 1). Also, the more firmly muscled and more
heavily ossified fishes, such as Hyperoglyphe, have greater swim-
bladder volumes than those with softer muscles and lighter bones,
such as Schedophiliis (Table 1 ). An exception is Nomeus, which
has relatively firm musculature but a small swimbladder.
The regression of the swimbladder is a gradual process, with
the sac diminishing and the gas gland becoming a small yellowish
mass before being completely resorbed. The mass representing
the regressed gas gland may persist in the mesentery beneath the
kidney for a considerable period of time after the swimbladder
becomes nonfunctional. In none of the stromateoids does the
swimbladder appear to become fat-filled upon regression as it
does in some deep-sea fishes (Marshall, 1960).
It is meaningful to consider the duration that the swimbladder
remains functional in the different stromateoid genera in rela-
tion to the time of change in habitat and mode of life. Nomeus
among the stromateoids appears to have the most intimate and
enduring association with jellyfishes, usually with the siphono-
phore, Physalia. The fish may remain with Physalia throughout
its life, although this is uncertain. Significantly, Nomeus retains
what seems to be a functional swimbladder longer than any other
stromateoid that has been examined. The largest specimen I have
studied (142.7 mm SL) had a relatively large sac with a some-
what contracted gas gland and was captured at the surface with
Physalia. It has not been possible to determine whether or not
the large specimens that have been found with Physalia were
mature.
The swimbladder of Peprilus triacanthus is usually completely
regressed by the time the fish reaches a length of 100 mm SL,
and this is about the size at which it has completely abandoned
jellyfish medusae. Large individuals (> 100 mm SL) of P. tria-
canthus do not hover as do the juveniles but swim continuously.
The pecioral fins increase in relative length with age and are used
more extensively for propulsion in adults than in juveniles. The
angle through which the pectorals are adducted apparently pro-
vides lift. (The locomotion and buoyancy of P. triacanthus are
being considered in a separate study.) Continuous swimming
with some degree of pectoral propulsion is probably the rule in
adult stromateoids.
In Stromateus fiatola, the swimbladder is greatly regressed at
a fish length of 75 mm SL, a size at which the fish seems to have
1970 SWIMBLADDER OF JUVENILE STROMATEOIDS 7
ended its association with medusae (Mansueti, 1963: 60). In
the size interval of 75 to 100 mm SL, this species loses the pelvic
fins, and its coloration changes from a vertically-banded pattern
to a more uniform one in which there are often dorsal spots.
Tetragoniirus is a strictly oceanic genus (Grey, 1955), and its
swimbladder is considerably regressed when the fish reaches a
size of 50 mm SL. The young have been found associated with
medusae in the surface waters (Mansueti, 1963: 60). According
to Haedrich ( 1967), the adults are probably members of the meso-
or bathypelagic fauna.
Finally, Pampus, a coastal genus and the one considered by
Haedrich (1967) to be the most advanced of the stromateids,
evidently has no swimbladder. Whether fishes of this genus as-
sociate with medusae as frequently as other stromateids is not
known. There are indications that they do not. According to
Suyehiro (1942), P. argenteus does feed to a certain extent on
jellyfishes in Japanese waters. However, studies by Kuthalingam
(1963) and Nath (1966) show that both juveniles and adults of
this species off the Indian coast are macroplankton feeders whose
diet largely depends upon seasonal changes in abundance of crus-
taceans and polychaetes; this may indicate that the young do
not regularly associate with medusae. Also, the pectoral fins of
Pampus become quite long early in life and do not greatly increase
in relative length with age as they do in Peprilus, another stro-
mateid, and certain other genera. At a fish size of 30 mm SL,
the pectoral length of Pampus argenteus is about 40 per cent of
the standard length compared to only about 30 per cent in Pep-
rilus paru, which has a very similar body shape. The pectoral
length in Peprilus triacanthus of the same size is about 25 per
cent of standard length. The relative length of the pectorals has
increased to 40 per cent in P. paru and to about 35 per cent in
P. triacanthus at a size of 80 mm SL. Thus, assuming that hover-
ing beneath objects and possession of a swimbladder are related
and that increased pectoral length is important in continuous swim-
ming, it seems that members of the genus Pampus acquire the adult
mode of locomotion and habit at an earlier stage than most other
stromateoids and, in so doing, completely dispense with the swim-
bladder.
A number of other fishes have regressed, age-dependent swim-
bladders. These include such shallow-water marine fishes as cer-
tain gobies, blennies, flatfishes, and most muraenid eels (N. B.
8 BREViORA No. 359
Marshall, personal communication ) and also some deep-sea fishes,
such as certain species of Cyclothone and Stomias, whose swim-
bladders, upon regressing, become invested with fat (Marshall,
1960). As in stromateoids, the regression of the organ in these
fishes is probably associated with habitat and/or internal changes.
Swimbladder regression is eventually to be the subject of a general
review.
Yet to be examined is the state of development of the swim-
bladder in newly-hatched stromateoid larvae. Since it is fully
formed in fishes as small as 9 mm SL, the organ must develop
quite early. The swimbladder may be of considerable importance
to larvae in orientation and in positioning the body for food-
capturing, although it is not known when the larvae begin to feed.
Indeed, the swimbladder, as I judge from its particularly early
regression in some stromateoids, may have its greatest functional
significance in the larvae and smallest juveniles.
ACKNOWLEDGMENTS
1 sincerely thank N. B. Marshall and Richard L. Haedrich for
reading and ofi'ering suggestions on the manuscript. I am also
grateful to Dr. Marshall for valuable advice on swimbladders
and for providing space and facilities in the British Museum (Na-
tural History). This work was supported by a NATO Postdoctoral
Fellowship awarded by the National Science Foundation.
LITERATURE CITED
Alexander, R. McN. 1966. Physical aspects of swimbladder function.
Biol. Rev., 41 (1): 141-176.
Fowler, H. W. 1936. The marine fishes of West Africa, based on the
collection of the American Museum Congo Expedition, 1909-1915.
Part 11. Bull. American Mus. Natur. Hist., 70(2): 607-1493.
Grey, M. 1955. The fishes of the genus Tetragoniirus Risso. DANA-
Report No. 41: 1-75.
Haedrich, R. L. 1967. The stromateoid fishes: systematics and a classi-
fication. Bull. Mus. Comp. Zool., 135(2): 31-139.
1969. A new family of aberrant stromateoid fishes from
the equatorial Indo-Pacific. DANA-Report No. 76: 1-14.
Haedrich, R. L., and M. H. Horn. 1969. A key to the stromateoid
fishes. WHOl Tech. Rept. No. 69-70, 46 pp. UNPUBLISHED
MANUSCRIPT.
1970 SWIMBLADDER OF JUVENILE STROMATEOIDS 9
Horn, M. H. 1970. Systematics and biology of the stromateid fishes of
the genus Peprilii.s. Bull. Mus. Comp. Zool., 140(5): 165-262.
Jordan. D. S., and B. W. Evermann. 1896. The fishes of North and
Middle America: a descriptive catalogue of the species of fish-like
vertebrates found in the waters of Ncrlh America north of the Isthmus
of Panama. Part 1. Bull. U. S. Nat. Mus., No. 47: 1-1234.
KuTHALiNGAM, M. D. K. 1963. Observations on the fishery and biology
of the silver pomfret, Pampiis argenteiis (Euphrasen), from the Bay
of Bengal. Indian J. Fish.. 10(1): 59-74.
Lane, C. E. 1960. The Portuguese man-of-war. Sci. Amer., 202(3):
158-168.
Mansueti, R. 1963. Symbiotic behavior between small fishes and jelly-
fishes, with new data on that between the stromateid, Pepriliis alepi-
dotus, and the scyphomedusa, Chrysaora qiiinqiiecirrha. Copeia, 1963
(1): 40-80.
Marshall, N. B. 1960. Swimbladder structure of deep-sea fishes in rela-
tion to their systematics and biology. Discovery Rept., 31: 1-122.
Maul, G. E. 1964. Observations on young live Miipus tnaculatus
(Giinther) and Miipiis oralis (Valenciennes). Copeia, 1964 (1):
93-97.
Nath, p. R. 1966. Biology and seasonal distribution of the pelagic food
fishes of Travancore coast. Kerala Univ. Pub., India, 1-140.
SuYEHiRO, Y. 1942. A study on the digestive system and feeding habits
of fish. Japan. J. Zool., 10 (1): 1-303.
BREVIORA
Muisenajnini of Comparative Zoology
Cambridge, Mass. 30 November, 1970 Number 360
MAMMALS FROM THE EARLY CENOZOIC OF
CHUBUT, ARGENTINA
George Gaylord Simpson
Abstract. Angelocabrenis daptes, new genus and species (Mammalia,
Marsupialia, Borhyaenidae), Coelostylodon florentinoameghinoi, new genus
and species (Mammalia, Notoungulata, ?Isotemnidae) and Coelostylodon
caroloameghinoi, new species, are described from the Casamayor forma-
tion, probably early Eocene. Knowledge of upper premolars of Didolodiis
(Mammalia, Condylarthra. Didolodontidae) is increased and the status of
Acoelodiis (Mammalia, Notoungulata, Acoelodidae) is discussed. The latter
genus and the family based on it are essentially indeterminate, and previous
usage of the names is unjustified.
INTRODUCTION
A visit to Mar del Plata, Provincia de Buenos Aires, Argentina,
early in 1970 enabled me for the second time to examine parts of
the important collections of fossil mammals in the Museo Munici-
pal de Ciencias Naturales of that municipality. In collections from
the Casamayoran Stage of Chubut, three specimens were found to
be of particular interest and to make especially important con-
tributions to knowledge. The Director of the Museo, Sr. Galileo
J. Scaglia, very kindly permitted me to study those specimens and
to publish descriptions and discussions of them, presented here-
with. I am again and increasingly indebted to Sr. Scaglia and to the
whole staff of the Museo for their courtesy and cooperation. The
accompanying illustrations were prepared by RaVae Marsh.
In the following, MMP precedes catalogue numbers of the
Museo Municipal de Ciencias Naturales de Mar del Plata and
MACN those of the Museo Argentino de Ciencias Naturales "Ber-
nardino Rivadavia," Buenos Aires.
While carrying out the research for this paper I was employed
jointly by the Museum of Comparative Zoology and the Univer-
sity of Arizona.
2 BREVIORA No. 360
Order Marsupialia lUiger
Family Borhyaenidae Ameghino
Angelocabrerus, new genus
Etymology. For the late Angel Cabrera, a great mammalogist,
who, among many other things, wrote an important study of bor-
hyaenids. This kind of nomenclature is Ameghinian and is con-
sonant with the related Arminiheringia. I have ventured to give
the compound an appropriately mascuUne ending.
Type-species. Angelocabrerus daptes, new species, infra.
Known distribution. Casamayoran, Argentine Patagonia.
Diagnosis. Specialized borhyaenines. Canines with closed,
rapidly tapering roots; short, heavy, fully enameled crowns. P.,
one-cusped, heavy, with small distinct talonid. M 1.4 essentially
two-cusped, with paraconid anterior and only slightly lingual to
the larger protoconid. No trace of metaconid. Talonids un-
basined, reduced to very slight, simple ledges. Protoconids and
paraconids truncated with wear on M^.g, becoming sharp points on
M4 with protoconid a high slender needle.
Discussion. As far as known, the dentition is similar to that
of the much later (Santacrucian) Borhyaena and by the same token
is also similar to the contemporary Casamayoran Arminiheringia.
The talonid reduction seems to have gone even further in Angelo-
cabrerus than in the other genera. The lower canine is unlike that
of Arminiheringia, with a shorter closed root and more fully
enameled crown. The way in which M^ wears, quite distinctive
from either Arminiheringia or Borhyaena, would seem to imply
different occlusion and hence different structure in the unknown
upper teeth, M^.^ , with which M^ occluded. This wear is much
as in Plesiofelis, considered by Cabrera (1927: 274-278) Des-
eadan in age and synonymous with Pharsophorus but almost cer-
tainly Mustersan and probably distinct from Pharsophorus. How-
ever, in Plesiofelis the molar talonids are considerably less re-
duced than in Angelocabrerus.
In Arminiheringia auceta, the only adequately known species
of its genus, there is a rapid increase in size of the molars pos-
teriorly, M4 being about twice as large as M^. The figures (some-
how omitted in Simpson, 1948) are here given in Table 1. It is
there shown that the increase is much less in Angelocabrerus dap-
tes, with M4 only about half again as large as M^. The increase
1970 EARLY CENOZOIC CHUBUT 3
is even less in Borhyaena. As far as the evidence goes, Angeloca-
hrerus could be ancestral to Borhyaena, and in that case the rate
of evolution in known parts must have been extremely slow.
Knowledge of the present genus is too incomplete, however, to
warrant a firm conclusion.
As in Arminiheringia and Borhyaena but to even more marked
degree, Pg is a large and heavy tooth. It here approximates M , in
dimensions. It has a single main cusp, with a long anterior and
short, nearly vertical posterior slope. There is a minute cuspule
at the anterior base. There is a distinct but small, shelflike talonid
with a single cuspule. This is absent in Borhyaena and also in the
only known specimen of Arminiheringia that might show it, but
the latter is so worn that a small talonid could have been present
originally. Except for the points already mentioned, the lower
molars are like Arminiheringia and Borhyaena in structure.
Angelocabrerus daptes, new species.
Etymology. Greek daptes, eater, gnawer, from the inferred
carnivorous, possibly ossifragous habits of the animals.
Holotype. MMP 967M, part of right mandibular ramus with
M2.4, left Pg, Ml, and M. probably of the same individual, two
lower canines somewhat broken, and small caniniform tooth and
tip of another doubtfully associated.
Hypodigm. Holotype only.
Horizon and locality. Casamayoran, south of Lago Colhue-
Huapi, Chubut, Argentina. The specimen was a surface find high
in the beds, and derivation from an overlying formation is pos-
sible but quite improbable.
Diagnosis. Only known species of the genus as diagnosed
above.
Discussion. The loose left M. has somewhat darker enamel
and is slightly less worn than the right M. in the mandibular frag-
ment. Its color and wear are more consonant with those of the
loose teeth identified as left Pg and M^. However, there can be
little serious doubt that those and the two loose lower canines do
in fact belong to the same individual as the mandibular fragment.
All were found together, they are congruent in size and structure,
and they add up to a unique specimen of a group extremely rare
in these beds. The two slender caniniform teeth are dubious and
I do question whether they belong to the same animal.
BREVIORA
No. 360
The diagnosis and discussion of the genus and the illustrations
make further description unnecessary.
Figure 1. Angelocabrerus daptes, new genus and species. Holotype,
MMP 967M. Right M2-4. A, buccal view. B, occlusal view. C, lingual
view. X 1.
Order Condylarthra Cope
Family Didolodontidae Scott
Didolodus sp. indet.
Specimen. MMP 696M, fragment of right maxilla with
2-4
Horizon and locality. Casamayoran of Caiiadon Vaca, tribu-
tary to the left (northwest) bank of the Rio Chico, Chubut, Ar-
gentina.
1970
EARLY CENOZOIC CHUBUT
Figure 2. Angelocabrerus daptes, new genus and species. Holotype,
MMP 967M. Left P^ - M,. A, lingual view. B, occlusal view. C, buccal
view. XI.
Figure 3. Angelocabrerus daptes, new genus and species. Holotype,
MMP 967M. Lower canine. X 1.
BREVIORA
No. 360
Discussion. This specimen is interesting because it shows the
coronal structure of P^'^ of Didolodus in relatively little worn
condition and because the proportions of these teeth are distinctive.
Comparison is mainly with MACN 10690, holotype of Didolo-
dus multicuspis Ameghino, the only other specimen of Didolodus
known to me that includes P^^. It is figured in Simpson, 1948,
text figures 25 and 26 and plate 10, figures 1 and 2. MACN 10738
includes P-, which has not been figured but was included in my
description of D. multicuspis (Simpson, 1948: 101). AMNH
2847 is a P'^ referred to D. minor by me (Simpson, 1948: 103)
but not separately described or figured.
As shown in Table 2, P- and P^ are each shorter than in the
holotype by 7 per cent, which does not in itself suggest specific dis-
tinction, and P^ has almost the same length in the two specimens.
However, all three teeth are more notably narrower in MMP
696M, by 16 per cent, 20 per cent, and 19 per cent for P-, P^,
and P^ respectively. A result is that all three teeth are longer
relative to their widths in MMP 696M. This is especially notice-
Figure 4. Didolodus sp. MMP 696M. Right PS-i, occlusal view. X 3.
able in P"^, which is distinctly transverse in MACN 10690 but
equidimensional in MMP 696M. P=^ of AMNH 28471, referred
to D. minor, is even more transverse than in the holotype of D.
multicuspis. Its width slightly exceeds that of P^ in MMP 696M,
although its length is decidedly (25 per cent) shorter. The in-
dividual represented by MMP 696M was probably closer to D.
multicuspis in over-all size, but the differences in some dimensions
and in proportions make reference doubtful either to that species
or to D. minor. At the same time, they do not warrant definition
of a new species, which in any case should preferably not be based
on upper premolars, for which there is so little comparative
material.
1970 EARLY CENOZOIC CHUBUT 7
Specimens of known origin referable to D. midticiispis without
much doubt are all from south of Lago Colhue-Huapi, and those
similarly referable to D. minor are from Canadon Vaca. MMP
696M is from Canadon Vaca but probably does not belong to D.
minor. Specimens and field data for Didolodus are still far from
sufficient for identification of populations within the genus and for
determination of their distribution.
P- of MMP 696M, unlike the less well-preserved specimens
previously known, is seen to have two distinct external cusps, con-
nate above the tips but still separated by grooves. These are at
least descriptively paracone and metacone, and the metacone is
only slightly lower and smaller than the paracone. The tooth is
completely surrounded by a cingulum, but this is feeble on the
middle of the labial face. The lingual slope from paracone plus
metacone to the labial cingulum is sUghtly uneven, but is without
really distinct cusps. P^ and P^ also have distinct paracone and
metacone but, unusually, the metacone is lower and smaller rela-
tive to the paracone progressively from P- to P^. P^ and P^ have
well-developed protocones, and the cingula do not cross their lin-
gual faces. A distinct cingular cusp or style is present on each
tooth anterior and slightly labial to the paracone and there is a
similar but smaller and less distinct cuspule posterior to the meta-
cone. Each tooth has a distinct protoconule but no metaconule.
There is no hypocone. It is also unusual that P^ is distinctly
shorter than P-^ although wider.
The structure of P- in this specimen seems to be rather different
from that in the holotype of D. multicuspis and more molariform.
Except for dimensions and proportions, apparent differences in
structure of P ^"^ are possibly due only to the more worn condition
of the holotype of D. multicuspis.
Although far from identical, there is considerable resemblance
between P^-^ of MMP 696M and the homologous teeth of North
American Phenacodus. A fairly close ancestral relationship is con-
firmed to that extent. However, considerable independent evolu-
tion is also suggested. For example, P^-^^ especially, of MMP
696M, are more distinctly molariform than their homologues in
Phenacodus.
8
BREVIORA
No. 360
Order Notoungulata Roth
Family Isotemnidae Ameghino
Coelostylodon, new genus
Etymology. Greek koilos, hollow, stylos, pillar, odon, tooth.
The name is meant to recall former reference to Acoelodiis and
resemblance to Pleurostylodon. It is also consonant with much
Ameghinian nomenclature.
Type-species. Coelostylodon florentinoameghinoi, new species,
injra.
Known distribution. Casamayoran, Argentine Patagonia.
Diagnosis. Primitive notoungulates with complete, nearly
closed dentition. Upper canine small and fully incisiform. Cheek
teeth brachydont, P--M^ soon wearing so that crown presents a
single fossa, without complex folds or anterior opening. M^-^
with flattened, slightly bifid lingual faces. M^^ with slight para-
style and paracone folds and very feeble metacone swelling on
ectolophs, no mesostyle. M^ subtriangular, with short but distinct
metaloph, longer than M' or M- and almost as long as broad.
Figure 5. Coelostylodon florentinoameghinoi, new genus and species.
Holotype, MMP 1723M. Right C and pa - M^. X W2.
Discussion. This genus is essentially that called Acoelodus
by Ameghino (1901: 467) and discussed by me (Simpson, 1967:
57) under that name, but that apphcation of the name can no
longer be sustained. The type-species of Acoelodus is A. oppositus
Ameghino, 1897 (p. 454). The holotype of that species is MACN
10770, a fragment of a left mandibular ramus with much worn
P 2.3 and part of Pj. That specimen is essentially indeterminate.
In 1901 Ameghino referred to the species a poorly preserved skull,
MACN 10753, and redefined the genus essentially on the basis of
1970 EARLY CENOZOIC — CHUBUT 9
that specimen. Desiring to validate as much as possible of Ame-
ghino's nomenclature, I accepted the reference and redefinition,
taking MACN 10753 as essentially a neotype in Ameghino's usage
and designating it as such (Simpson, 1967: 58). However, that
action is invalid under the present code of nomenclature (Stoll
et al., 1964, Article 75) because in fact the holotype, MACN
10770, has not been lost or destroyed.
If now there were a reasonable probability that the referred skull
and the holotype belong to the same genus and species, the type
designation would have no practical importance and one could
continue to use the name Acoelodus for the genus and A. oppositus
for the species represented by the referred skull. In fact, however,
as I already mentioned in previous discussion (1967), there is no
good evidence that the two specimens are of the same genus and
species, and there is some contrary evidence. The contrary evi-
dence is weightier than I previously indicated. If the comparative
sizes of P 2.3 and P^-^ jn Acoelodus were approximately as in
Pleurostylodon, a reasonable assumption, then P ^-s of the skull
MACN 10753 are some 22 to 44 per cent larger in various dimen-
sions than would be expected from the holotype of Acoelodus
oppositus and are also different in relative sizes and proportions.
It is thus highly improbable that the two specimens are conspecific,
and if they are not conspecific, there is no reason to consider them
congeneric.
The genus represented by the holotype of Acoelodus oppositus
is indeterminate, a notoungulate incertae sedis as far as my knowl-
edge and judgment go. The genus represented by MACN 10753
is determinate and is distinct from any other for which there is
comparable material known to me. This conclusion is reinforced
by discovery of another specimen, MMP 723M, that can be re-
ferred to the same genus as MACN 10753 but is specifically dis-
tinct. In order to avoid possible further confusion with Acoelodus,
the species represented by MMP 723M is made type-species for
the new generic name.
In order to avoid still another confusion, it must be noted that
none of Ameghino's figures labeled "Acoelodus oppositus" in sev-
eral of his publications are conspecific or congeneric either with
the holotype of that species or with MACN 10753 (for details and
discussion see Simpson, 1967: 58-59).
Ameghino considered Acoelodus as closely related to Oldfield-
thomasia and so placed the latter in his family Acoelodidae, but
10 BREVIORA No. 360
that was based on specimens dubiously or incorrectly referred to
Acoelodus, including at least one that in fact belongs in Oldfield-
thomasia. Since the holotype of the type-species of Acoelodus is
not identifiable as to family, the name Acoelodidae has no estab-
lished significance. For that reason, I named a family Oldfield-
thomasiidae for Oldfieldthomasia and its probable relatives (Simp-
son, 1945: 126). In revision of the Casamayoran fauna, I put
MACN 10753, under the incorrect reference name Acoelodus
oppositus, in the Oldfieldthomasiidae. The genus Coelo stylo don,
to which that specimen is now referred as holotype of C. caro-
loameghinoi, is of uncertain family position. It differs from all
adequately characterized previously named genera of both the
Oldfieldthomasiidae and the Isotemnidae, but has resemblances to
both families. Present reference to the Isotemnidae is very tentative.
The upper molar structure is most nearly similar to that of Pleu-
rostylodon, an isotemnid, among adequately known genera, but
Coelostylodon differs from Pleurostylodon and other isotemnids in
its small, incisiform canine and various other details. The canine
is more like that of Oldfieldthomasia, but the molars are quite
different.
Coelostylodon florentinoameghinoi , new species
Etymology. For Florentino Ameghino, famed describer of
most of the Casamayoran fauna. Combination of given and family
names is a nomenclatural device that he often used.
Holotype. MMP 723M, nearly complete but badly crushed
skull.
Hypodigm. The holotype only.
Horizon and locality. Lowest Casamayoran beds in the bar-
ranca south of Lago Colhue-Huapi, Chubut.
Diagnosis. Significantly larger than C. caroloameghinoi (see
Table 3 ) . Posterolabial angle of M^ less projecting.
Discussion. The teeth are deeply worn in both of the holo-
types now referred to this genus. As preserved, there is no
marked, discernible difference in structure of the cheek teeth.
MMP 723M clearly has the small canine considered characteristic
of the genus. P^^^are almost completely fragmented, but seem to
have been quite small, perhaps more so, relatively, than in MACN
10753. The posterolabial corner of M^ does not project so dis-
tinctly as in MACN 10753.
1970 EARLY CENOZOIC CHUBUT 1 1
The skull is too badly crushed to make out much significant de-
tail. It seems to have been a primitive, unspecialized notoungulatc
skull generally similar to that of Pleurostylodon.
Coelostylodon caroloameghinoi , new species.
Acoelodus oppositus, in error, Ameghino, 1901: 365; Simpson,
1967: 58 and plate 11, fig. 1.
Etymology. For Carlos Ameghino, who discovered the Casa-
mayoran fauna and found the holotype of this species.
Holotype. MACN 10753, poorly preserved anterior part of
skull.
Hypodigm. The holotype only.
Horizon and locality. Casamayoran, barranca south of Lago
Colhue-Huapi.
Diagnosis. Significantly smaller than C. florentinoameghinoi
(see Table 3). Posterolabial angle of M^ sharply produced
posteriorly (or distally).
Discussion. In 1901 and thereafter Ameghino based his con-
cept of Acoelodus oppositus mainly on this specimen, but he did
not figure it, and none of the specimens figured by him as Acoe-
lodus oppositus belong to the present genus or species. The holo-
type is adequately figured in my previous memoir, as cited above.
REFERENCES
Ameghino, F. 1897. Mammiferes cretaces de I'Argentine. Deuxieme
contribution a la connaissance de la fauna mammalogique des couches
a Pyrotherium. Inst. Geog. Argentina, 18: 406-429, 431-521.
. 1901. Notices preliminaires sur des ongules des terrains
cretaces de Patagonia. Bol. Acad. Nac. Cien. Cordoba, 16: 350-426.
Cabrera, A. 1927. Datos para el conocimiento de los dasiuroideos fosiles
argentinos. Rev. Mus. La Plata, 30: 271-315.
Simpson, G. G. 1945. The principles of classification and a classification
of mammals. Bull. American Mus. Nat. Hist., 85: i-xvi, 1-350.
. 1948. The beginning of the Age of Mammals in South
America. Part 1. Bull. American Mus. Nat. Hist., 91: 1-232.
. 1967. The beginning of the Age of Mammals in South
America. Part 2. Bull. American Mus. Nat. Hist., 137: 1-259.
Sinclair, W. J. 1906. Marsupialia of the Santa Cruz beds. Repts. Prince-
ton Univ. Exped. Patagonia, 1896-1899, vol. IV, part III: 333-460.
Stole, N. R., et al. 1964. International code of zoological nomenclature
adopted by the XV International Congress of Zoology. International
Trust for Zoological Nomenclature, London.
12 BREVIORA No. 360
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1970
EARLY CENOZOIC CHUBUT
13
TABLE 2
Measurements in millimeters of upper premolars of Didolodus.
p2
p3
p4
Length
Width
L/W
Length
Width
L/W
Length
Width
L/W
D. luulticuspis
MACN 10690
7.6
7.4
1.03
7.5
9.5
.79
7.0
10.0
.70
D. minor
AMNH 28471
5.7
7.8
.58
D. sp.
MMP696M
7.1
6.2
1.15
7.6
7.6
1.00
6.5
8.1
.80
TABLE 3
Comparative measurements in millimeters of dentitions of holo-
types of Coelostylodon florentinoameghinoi and C. carolo-
ameghinoi.
p4
Ml
M2
M3
Length
Width
Length
Width
Length
Width
Length
Width
C florentinoameghinoi
C caroloameghinoi
MMP723M
MACN 10753
9.9
7.5
12.5
9.9
Ca.lO
8.0
15.4
11.5
12.6
8.5
17.0
11.8
15.1
10.0
15.4
10.7
BREVIORA
Mnaseiam of Compsirative Zoology
Cambridge. Mass. 30 November, 1970 Number 361
ADDITIONS TO KNOWLEDGE OF THE ARGYROLAGIDAE
(MAMMALIA, MARSUPIALIA) FROM THE LATE
CENOZOIC OF ARGENTINA
GEORGE GAYLORD SIMPSON
Abstract. A recent monograph of the Argyrolagidae is supplemented
by observations on specimens not previously seen by the author. The
holotype of Argyrolagiis palineri confirms previously published data. An
additional specimen of Argyrolagiis scagliai adds to i<nowledge of the
coronal pattern of lower molars. Argyrolagiis parodii is a hitherto dubious
species the holotype of which was destroyed. A second specimen is made
neotype; it validates the species and permits redefinition.
A monograph of the Argyrolagidae (Simpson, 1970) was com-
pleted early in 1968 and an addendum included late in that year.
Early in 1970 a visit to Argentina made possible the study of
several specimens not available or not known to me when the
monograph and addendum were written. These add significantly
to knowledge of the family.
For the facilities for making and the privilege of publishing
these observations I am greatly indebted to the authorities of the
Museo de La Plata and especially to Dr. Rosendo Pascual and
Sr. Jorge Zetti. The accompanying photographs were made under
the direction of Sr. Zetti and provided for this publication by the
Museo. In Buenos Aires Sr. Guillermo del Corro courteously
made available the type of Argyrolagiis palmeri. The drawing was
made by RaVae Marsh from a sketch by me. While making this
study I was employed jointly by Harvard University and the Uni-
versity of Arizona.
In the following all measurements are in miUimeters: L r=z length
(anteroposterior or mesiodistal). W = width (bucco-lingual).
Ljj = length of trigonid. Lp = length of talonid. W^ =r width of
trigonid. Wj, = width of talonid. MACN = Museo Argentino de
2 BREVIORA No. 361
Ciencias Naturales "Bernardino Rivadavia." MLP ■=. Facultad
de Ciencias Naturales y Museo de La Plata. MM? = Museo
Municipal de Ciencias Naturales de Mar del Plata.
Family Argyrolagidae Ameghino, 1904
Genus Argyrolagus Ameghino, 1904
Argyrolagus palmeri Ameghino, 1904
Holotype. MACN A53-4, part of left mandibular ramus, ori-
ginally with I J and M ^.4, collected by Carlos Ameghino at Monte
Hermoso in 1904.
Remarks. For other formal data see Simpson, 1970: 11.
When that account was written I had not seen the type, which is
in the Ameghino Collection in Buenos Aires and catalogued as
above. I^ has been lost since the specimen was figured by F.
Ameghino (1906, fig. 221) and L. Kraglievich (1931, fig. 2).
Both of those figures represent the specimen accurately. New
measurements on the specimen agree reasonably well with those
taken from the earlier figures (given in Simpson, 1970, Table 1 ),
see present Table 1.
Argyrolagus scagliai Simpson, 1970
Addition to hypodigni. MPL 49-1X-7-1, right mandibular la-
mus with alveoli of I ,0 and P. and all of Mi,. "Miramar, prov.
de Bs. Aires," probably Chapadmalal formation. Presented to
the MLP by the widow of Dr. Santiago Roth in August, 1924.
Description. For formal data on the species see Simpson,
1970: 12. This specimen has M^.^ beautifully preserved and less
worn than in others known. In this genus the hypselodont molars
rapidly lose the coronal pattern by wear and become columns with
characteristic outline but no internal structure. Here even Mj,
although somewhat worn, preserves some coronal features. M^
is also fully erupted and beginning to wear. The teeth do not
taper in the alveoli and would not increase in size with wear. The
dimensions of the exposed ends of these young teeth are com-
parable with those of old individuals of the species (see Table 2).
All the molars retain traces of a fossettid in the trigonid. M, has
a somewhat more complex outline than in other, individually older
specimens considered to be of the same species (see Fig. 2).
Most of the crown is horizontally truncated by wear, but there is
a steeply sloping anterolingual wear facet. On the anterior face
1970
ARGYROLAGIDAE
there is a well-developed cingulum, not yet worn, enclosing a small
pocket. The preservation makes possible separate trigonid and
talonid measurements in Table 3.
A.
B.
C.
Figure 1. Argyrolagus scaglicii Simpson, 1970. M.L.P. No. 49-1X-7-1.
right mandibular ramus with M ^,^ . A, buccal view. B. occlusal view.
C, lingual view. Approximately X 4i/4.
BREVIORA No. 361
Figure 2. Argyrolagus scagliai Simpson, 1970. M.L.P. No.49-IX-21,
right M^. Occlusal view, diagrammatic sketch. Approximately X 10.
Argyrolagus parodii Rusconi, 1933
Neotype. MLP 62-V11-27-81, right mandibular ramus with
I^, alveolus of I., and PfM,, from the Playa las Palomas, near
the Balneario Chapadmalal. Collected by the personnel of MLP
in August, 1961.
Revised diagnosis. Smaller than A. palmeri or A. scagliai.
Trigonids somewhat more triangular, paraconids medial. Talonids
relatively short. Labial and lingual sulci directly opposite. Talonid
of Mj somewhat reduced, but decidedly larger than in Microtra-
gulus. Buccal extension of coronoid crest strong.
Discussion. In a letter written shortly before his lamented
death, Sr. Rusconi informed me that the holotype of this species,
in his private collection, had been destroyed. The available figures
and description of that holotype are somewhat equivocal, as dis-
cussed in Simpson, 1970: 14-15. There is little doubt, however,
that MLP 62-Vn-27-81 does belong to the same species. It is
very close to the size indicated by Rusconi in text and figures, as
here shown in Table 4, and it also shares the most likely morpho-
logical distinctions of the holotype. The present specimen can
be confidently referred to Argyrolagus and it shows that A. parodii
is distinct from any other known species — both points left in
some doubt by the destruction of the holotype. This case meets
all the conditions for designation of a neotype set forth in Article
75(a) of the International Code of Zoological Nomenclature
adopted by the XV International Congress of Zoology, and the
present treatment supplies all the qualifying conditions specified in
Article 75(c) of the Code.
The locality where the holotype was found is not precisely
known but was somewhere along the shore four or five kilometers
1970
ARGYROLAGIDAE
Figure 3. Argyrolagiis parodii Rusconi, 1933. Neotype, M.L.P. No.
62-V!!-27-81. right mandibular ramus with I^ and P.^ - M,. A, buccal
view. B, occlusal view. C, lingual view. Approximately X 6.
BREVIORA
No. 361
northeast of Miramar, Buenos Aires Province. The neotype is
from the same sequence of exposures on the shore about llVz
to 13V2 kilometers farther northeast. The holotype was probably
from the Chapadmalal formation, as restricted, but possibly
Vorohue. (See J. L. Kraglievich, 1952.) There is no clearly
/ '-*'S
n
^
'I
,i.r
I""*'
A.
B.
c.
D.
Figure 4. Argyrolagidae, genus and species undetermined. M.L.P. No.
59-IX-28-98, right tibiofibula lacking unfused portion of fibula. A, lateral
view. B, anterior view. C, medial view. D, posterior view. Approximately
X 2.
1970 ARGYROLAGIDAE 7
established difference between the faunas of the two formations.
The neotype is almost certainly from the restricted Chapadmalal
formation. The two are probably of the same age, and in any
case can hardly have a faunally appreciable difference in age.
The specimen now made neotype of Argyrolagus parodii is the
one figured, without catalog reference, as Microtragulus argentinus
by Ringuelet (1966, plate X, figs. I, J) and used as a basis {Ibid.,
pp. 58-59) for redefinition of the genus Microtragulus (taken as
a synonym of Argyrolagus) and the family Microtragulidae
(equated with Argyrolagidae). As previously pointed out (Simp-
son, 1970: 65-67), the redefinitions are valid for the specimen in
question but are too restricted for the genus and family as wholes.
The preceding diagnosis and accompanying measurements
(Table 4) and photographs (Fig. 3) make detailed description
unnecessary.
Argyrolagidae gen. et sp. indet.
MLP 59-IX-28-98 is a tibiofibula lacking the free part of the
fibula, collected by Dr. J. Frenguelli from the "Chapadmalense"
of Baliza Chica, northeast of Miramar. At this locality it would be
either from a very high level in the Chapadmalal formation, re-
stricted, or from the overlying Vorohue formation. Its length is
53.5 mm, compared with 60.4 mm for MMP 7855, holotype of
Argyrolagus scagliai. It also differs from the latter in that the
proximal part of the cnemial crest ends abruptly slightly below
rather than at the articular surfaces and also ends abruptly distally
rather than passing gradually into the shaft. This specimen may
belong to Microtragulus reigi, but association cannot now be estab-
lished.
REFERENCES
Ameghino, F. 1906. Les formations sedimentaires du Cretace superieur
et du Tertiaire de Patagonie. Ann. Mus. Nac. Buenos Aires, ser. 3, 8:
1-568. Also published, with Spanish translation, as volume 16 of the
Obras (Ameghino, 1913-1936).
Kraglievich, L. 1931. Cuatro notas paleontologicas, sobre Octomylodon
aversus Amegh., Argyrolagus palmeri Amegh., Tetrastyhis montanus
Amegh., y Munizia paranensis. Rev. Soc. Argentina Cien. Nat., 10:
22-266. Also in the Obras (Kraglievich, 1940: 581-602).
8 BREVIORA No. 361
Kraglievich, J. L. 1952. El perfil geologico de Chapadmalal y Miramar,
Provincia de Buenos Aires. Rev. Mus. Municipal Cien. Nat. y. Tradic.
Mar del Plata, 1 : 8-37.
RiNGUELET, A. B. DE. 1953. Marsupialia. In A. V. Borello, ed., Paleon-
tografia Bonaerense, Comis. Invest. Cien.. Prov. Buenos Aires, La Plata,
fascicule IV: 46-59.
RuscoNi, C. 1933. New Pliocene remains of diprotodont marsupials from
Argentina. Jour. Mammal., 14: 244-250.
__. 1936. La supuesta afinidad de Argyrolagiis con los Typotheria.
Bol. Acad. Nac. Cin. Cordoba, 33: 173-182.
Simpson, G. G. 1970. The Argyrolagidae, extinct South American Mar-
supials. Bull. Mus. Comp. Zool., 139: 1-86.
1970 ARGYROLAGIDAE 9
TABLE 1
Measurements of teeth of holotype of Ar^yrola^us palmeri
M, M,. M3 M4 LMu4
L W L W L VV L W
MACN A53-4 1.9 1.1 2.1 1.3 2.0 1.3 2.0 1.1 7.8
TABLE 2
Comparative measurements of teeth of Argyrolagus scagUai
M, M2 Ma M4 LMi^
L W L W L W L VV
Holotvpe, MMP 7855 1.7 1.4 2.0 1.7 2.0 1.6 2.2 1.2 8.5
MLP49. IX. 7.1 1.8 1.7 2.3 1.9 2.3 1.8 2.2 1.4 8.6
TABLE 3
Trigonid and talonid measurements of Argyrolagus scagliai, MLP
49.IX.7.1.
Ml M-. Ms M4
La Lp Wa Wp La Lp W^ Wp L^ Lp Wa Wp L^ Lp W^ Wp
1.2 0.6 1.6 1.7 1.4 0.9 1.9 1.9 1.5 0.8 1.8 1.6 1.3 0.9 1.4 1.3
TABLE 4
Measurements of destroyed holotype of Argyrolagus parodii, from
text and figures of Rusconi, and of neotype MLP 62-VII.27.81.
Holotype:
Rusconi, 1933, te.xt
Rusconi, 1933, fig. la
Rusconi, 1936, fig. 12 —
Neotype 1.6 1.4 1.8 1.5 1.7 1.4 1.7 1.1 6.7
M,
M-.
M3
A
^4
LM,
L W
L \V
L W
L
\v
— — ■
— —
1.9 —
1.3
—
—
— —
— —
1.8 1.6
1.6
1.2
—
— —
— —
1.6 1.4
1.6
1.2
—
BREVIORA
Mmseiuiiiii of Compsirative Zoology
Cambridge, Mass. 30 November, 1970 Number 362
ADDITION TO KNOWLEDGE OF GROEBERIA
(MAMMALIA, MARSUPIALIA) FROM THE
MID-CENOZOIC OF ARGENTINA
George Gaylord Simpson
Abstract. Groeberia pattersoni, new species, from the Divisadero
Largo formation, is based on the second known specimen of that genus.
Some information on the hitherto unknown skull of genus and family is
provided. Reference to the Marsupialia is supported, but affinities within
the Marsupialia remain dubious.
The extraordinary fossil marsupial genus Groeberia has hitherto
been known from a single specimen, a fragmentary mandible,
named and described by Patterson (1952). A second specimen
was found by Dr. Edgardo Rolleri of the Yacimientos Petroliferos
Fiscales (Argentine government petroleum bureau) and deposited
in 1968 in the Museo de La Plata. Dr. Rosendo Pascual kindly
referred it to me for study, which I carried out in La Plata early
in 1970, and for publication, here presented. Sr. Jorge Zetti, as-
sistant to Dr. Pascual, facilitated the study and arranged for photo-
graphs, provided by the Museo for this publication. The accom-
panying drawings were made by RaVae Marsh on the basis of
sketches by me. I am also indebted to Dr. A. J. Amos, Dean of
the Faculty of Natural Sciences and the Museum of La Plata, and
to Sr. G. J. Scaglia, Director of the Museo Municipal de Ciencias
Naturales of Mar del Plata, who sent the holotype of G. minoprioi
on loan to La Plata for direct comparison with the present speci-
men.
The research for this paper was performed while I was employed
jointly by the University of Arizona and the Museum of Com-
parative Zoology.
2 BREVioRA No. 362
Class Mammalia Linnaeus, 1758
Order Marsupialia Illiger, 1811
Family Groeberiidae Patterson, 1952
Genus Groeberia Patterson, 1952
Groeberia pattersoni, new species
Etymology. For Bryan Patterson, who named and described
the genus and its type-species, G. ininoprioi.
Holotype. Museo de La Plata No. 68-VI-27-1, partial skull
and mandible.
Hypodigm. Holotype only.
Horizon and locality. Divisadero Largo formation, [in the
general vicinity of] Mina Atala, Mendoza, Argentina.
Diagnosis. Anterosuperior part of symphysis much more
slender and incisors smaller than in G. minoprioi. M 3.4 also, but
less, smaller. See Table 1.
Identification. The holotype of the type-species Groeberia
minoprioi includes most of the mandibular symphysis and incisors,
left Mo. .J , and broken bases of M, and M, (see Patterson, 1952).
The present specimen also preserves most of the symphysis and
lower incisors and has the broken bases of left M 3.4. These parts
are morphologically closely similar in the two specimens and quite
unlike any other animal known to me. Reference to the same genus
is indicated. Both are from the Divisadero Largo formation and
'&'-
TABLE 1
Comparative measurements in millimeters of holotypes of
Groeberia minoprioi and G. pattersoni.
G. minoprioi
G. pattersoni
Transverse at narrowest
part of symphysis, at
postincisive diastema
ca. 5.8
ca. 3.4
Transverse, across both
incisors at alveoli
ca. 5.8
ca. 3.4
M3 Length
2.3
ca. 1.8
M., Width
1.6
ca. 1.4
M^ Length
ca. 2.3
ca. 1.5
M^ Width
ca. 1.8
ca. 1.3
Measurements of M.. of G. pattersoni and of M^ of both speci-
mens are on broken bases of teeth and are rough approximations.
1970 GROEBERIA 3
near the same locality: the holotype of G. minoprioi from one-half
kilometer east of the Cerro Divisadero Largo (Patterson, 1952:
3) and that of G. pattersoni recorded as "Mina Atala," which
would be about IVi kilometers northeast of the previous locality
but which must be a rough approximation, as the Mina Atala is
not on the Divisadero Largo formation.
The holotype of G. minoprioi is from Minoprio's stratum F
(see map and stratigraphic discussion in Simpson, Minoprio, and
Patterson, 1962). The level of the present specimen is not so
precisely known. Although no faunal change has been demon-
strated within the formation, its deposition may have covered a
considerable span of time and the two specimens may not be very
closely contemporaneous. There is some presumption that speci-
mens so similar in morphology and provenience are conspecific,
but, as far as I know, the difference in slenderness of the sym-
physis and sizes of the teeth are greater than within adults of
any one species of marsupials. This difference can hardly be due
to greater age of the holotype of C. minoprioi. The incisors of the
holotype of G. pattersoni do not taper in the alveoli, hence they
could age considerably without increasing in diameter at the alveoh.
Although measurements are imprecise, the length of M^ is about
50 per cent longer in the holotype of G. minoprioi, and these
brachydont teeth do not grow after eruption. The holotype of G.
pattersoni evidently had all teeth erupted and some skull sutures
closed, as in fully adult animals. Thus specific separation is in-
dicated.
Age. Simpson, Minoprio, and Patterson (1962: 290) con-
cluded that "the age of the Divisadero Largo fauna is approxi-
mately early Deseadan or latest pre-Deseadan," but that knowledge
at that time did not warrant basing on it the apparently missing
mammalian age-stage between Mustersan and Deseadan. How-
ever, on evidence not fully stated, Pascual, Ortega, Gondar, and
Tonni (1965) proposed a "Divisaderense" (in English, Divisa-
deran) mammal age-stage as intermediate between Mustersan and
Deseadan and separated from each by a hiatus. They tentatively
correlated it with the Ludian of Europe and Duchesnian of North
America as latest Eocene. As those authors also did recosnize.
the data do not really permit close correlation, and I believe that
all one can now say is that the age in terms of the European epochs
may be somewhere around late Eocene or early Oligocene, hence,
in terms of absolute age, more or less middle Cenozoic.
4 BREVIORA No. 362
Description. As found, the specimen included at least the
anterior part of the skull and most of the mandible in articulation.
Some time before the specimen came into the control of the Museo
de La Plata, the skull and mandible were separated and both were
severely damaged. Nevertheless, what remains adds considerably
to knowledge of this remarkable and enigmatic genus.
The two lower incisors are preserved for a length of about 12
mm within their alveoli. The anterior ends are broken and the
posterior ends have been ground smooth, presenting the appear-
ance seen in Figure IB. The pulp cavity is here open, and there
is no sign of root formation, so these teeth were clearly hypselo-
dont/ as in G. minoprioi. The teeth curve so that the posterior
parts were nearly horizontal, in an odd medial posterior projection
of the symphsis, and the extra-alveolar parts would have been
nearly vertical. Enamel is lacking on the medial and dorsal faces,
and at this depth in the alveoli even the dentine has not quite
closed the pulp cavity dorsally. Heavy enamel occurs ventrally,
and this extends, thinning as it goes, onto the lateral faces, more
so than in the extra-alveolar parts of the incisors of the holotype
of G. minoprioi.
There is a short diastema posterior to the lower incisors and
then a series of cheek teeth, probably four as in G. minoprioi, al-
though here they cannot be definitely counted. The bases of what
are almost certainly the last two cheek teeth can be made out, but
the crowns are not visible. These teeth are brachydont as in G.
minoprioi and similar in outline but perhaps slightly less elongate.
There are two pairs of upper incisors, here designated I' and 1-
for convenience, although their ancestral homologies are unknown.
1 Dictionaries give "hypselodont" as a variant of "hypsodont," and the
most recent authoritative odontology, Peyer (1968), uses "hypselodont"
in place of "hypsodont." It is, however, more convenient to adopt a dis-
tinction sometimes made by mammaiogists, especially paleomammalogists.
I define as hypsodont a tooth that eventually develops one or more roots
but that has a crown definitely higher than those roots or than any of its
horizontal dimensions and, as hypselodont, a tooth that never forms a
root but continues to grow and to extrude new parts from the alveolus
throughout life. Peyer was not a mammalogist and devoted relatively
little attention to mammal teeth. Other odontologists have often oriented
their work on groups, especially Homo, in which hypselodont teeth (in
my sense) do not occur. The distinction is of great functional importance.
1970
GROEBERIA
B.
bone
enamel
dentine
matrix in
pulp cavity
Figure I. Grocheria pattersoni, new species. Holotype, M.L.P. No.
68-VI-27-1. A, sketch and measurements of wear surface of P, approxi-
mately X 6. B, sketch of posterior (intra-alveolar) exposure of paired
lower incisors, approximately X 5.
P is a large, strongly curved tooth with an alveolus extending
posterodorsally far back in the facial region to above the infra-
orbital foramen. It and I- are probably both hypselodont. The
cross section is peculiar, with an oblique, long, slightly convex,
buccal, enamel-covered face; a flat, anteroposterior, enamelless,
anteromedial face; and a likewise enamelless, concave, postero-
medial face. The three faces meet at definite angles, approximately
right angles at both ends of the anteromedial face but strongly
acute between the buccal and posteromedial faces. (See Fig. lA.)
The enamel is nearly smooth but with slight wavy longitudinal rib-
bing. Right and left P are close to each other on their medial
faces.
I- is much smaller than P, measuring about 1.1 mm across the
buccal face as compared with about 2.5 mm for P. I- is also
enameled on the buccal face, and probably not elsewhere. The
cross section cannot be clearly determined as the specimen is
preserved. I- is less curved longitudinally than P and although
BREVIORA
No. 362
the extra-alveolar parts of the two are in contact, the alveoli di-
verge. In the most probable orientation of the skull, the extruded
part of P is slightly recumbent and that of I- is nearly vertical.
Figure 2. Groeheria pattersoni. new species. Hololype. M.L.P. No.
68-VI-27-1. Fragment of symphysis with parts of lower incisors. A,
supero-pcsterior or lingual view. B, infero-anterior or genial view. C, right
lateral view (in more anatomical orientation the anterior parts of the in-
cisors would be nearly vertical). Approximately X 4.
1970
C.ROEBERIA
'mx-mx
suture?
IFO
foramen ?
Buccal
to M^
Mx- Ju
suture ?
Figure 3. Groehcria pcittersoni. new species. Holotype. M.L.P. No.
68-VI-27-1. Anterior part of skull, left lateral view, photograph and ex-
planatory sketch. "Buccal to M-^" indicates a point on matrix, formerly
covered by the zygoma, medial to which is a broken cheek tooth identi-
fied as probably M'\ IFO = infraorbital. Mx-Ju = maxillo-jugal. NA
= nasal. OR = orbit. Pmx-mx = premaxillo-maxillary. Approximately
X 4.
8
BREVIORA
No. 362
IFO forame
HR
Mx-Ju
suture ?
Figure 4. Groeberia pattersoni, new species. Holotype. M.L.P. No.
68-VI-27-1. Anterior part of skull, right lateral view, photograph and
explanatory sketch. CO = fragment of coronoid process of mandible. DPZ
= descending plate of zygoma. HR = fragment of horizontal ramus of
mandible. IFO = infraorbital. Mx-Ju i=: maxillo-jugal. OR = orbit.
Approximately X 4.
1 970
GROEBERIA
4"';
'^
•^i
B.
Figure 5. Groeheria pattersoni, new species. Holotype M.L.P. No.
68-VI-27-1. Anterior part of slcull. A, dorsal view. B. palatal view. Ap-
proximately X 4.
10
BREVIORA
No. 362
There is a diastema posterior to I- and then cheek teeth, but noth-
ing definite can be made out for the latter, except that they are
small and brachydont.
The preorbital part of the skull is notably short and deep. In
the most probable orientation, it (or the snout) is convex dor-
sally as a whole and curves downward anteriorly. The nasals are
accordingly downcurved and end shortly anterior to P. The infra-
orbital foramen is not definitely visible as the specimen is pre-
served, but must be of moderate size and in a normal position on
the face anteroventral to the orbit and about half way between it
and 1-, where there is a depression obscured by matrix on the
specimen. On the left side, what is probably part of the pre-
maxillo-maxillary suture is visible just anterior to that depression.
The orbit is rather small and relatively anterior in position, its
anterior rim probably in advance of the cheek teeth and certainly
well in advance of what is identified as M-\ The stout root of the
zygoma below the orbit had an expanded suborbital process or
plate, the full extent of which cannot be determined. On both
sides, a possible but uncertain maxillo-jugal suture can be seen
Figure 6. Groeberia pattersoni, new species. Holotype, M.L.F. No.
68-VI-27-1. Anterior part of skull, anterior view. Approximately X 4.
1970 GROEBERIA I 1
below the ventral border of the orbit. Posterior to this, and hence
on the jugal if the possible suture is such, is a small ventral postor-
bital process. Uncertainly but probably, there was no dorsal post-
orbital process, and the orbit was therefore probably open.
The palatal surface is both poorly exposed and poorly preserved,
so that little can be made out there with sufficient probability.
However, its dorsal (intranasal) surface is partly exposed, and
some details can be made out in cross section at the broken pos-
terior surface of the specimen. A palatal vacuity was probably
absent or small if present. It appears that the palate between the
cheek teeth was deeply arched (concave ventrally). The coro-
noid process of the mandible can be seen lateral to a posterior
tooth, perhaps M'' or M^, and lateral to that a section of a deep
(vertically) but thin (laterally) zygoma. (See diagram, Fig. 7.)
palate {^probable
zygoma \^ K^ cartilage
mandible
Figure 7. Groeberia pattersoni, new species. Holotype, M.L.P. No.
68-VI-27-1. Rough diagram of section at posterior break of specimen as
preserved. Not to scale.
Affinities. This specimen adds considerably to data bearing
on affinities of the genus, without settling the matter. Patterson's
(1952: 3) judgment that Groeberia should be distinguished at the
family level, as Groeberiidae, is strongly confirmed.
Patterson's evidence that Groeberia is a marsupial was in small
part positive, largely negative, and partly indirect. The present
specimen adds no positive evidence, strengthens the negative evi-
dence, and does not change the indirect evidence. The only
really positive evidence from the holotype of G. ininoprioi for
marsupial affinities in general (as distinct from caenolestoid affin-
ities in particular, see below) is the probably inflected angle of
the mandible. Even this is not quite certain, because the angle
itself is not known, but a crest that would have led to it does
decidedly suggest inflection. An inflected angle is strong, but not
conclusive, evidence, as a few marsupials do not have the angle
inflected and a few placentals do. G. minoprioi may have had four
12 BREVIORA No. 362
lower molars, and that would again be strong but not fully con-
clusive evidence, but the possibility that its cheek teeth include
one premolar and three molars is not entirely excluded. The
known parts of the skull of C pattersoni have no evident features
strongly characteristic either of marsupials or of placentals. The
absence of palatal vacuities would be more like most placentals,
but it is uncertain and there are a number of marsupials without
such vacuities.
The negative evidence is that Groeberia has no known features
that would make reference to the Marsupia'ia impossible or highly
improbable but does have known features that make reference
to any other order highly improbable. Here the new specimen
confirms and adds somewhat. The habitus is more or less rodent-
like, but the two upper incisors are unlike those of any rodent.
They are somewhat like those of lagomorphs, but the cheek teeth,
even what little can be seen of them in this specimen, definitely
rule out pertinence to that group. Limited resemblances of the
mandible to those of certain primates, such as the early Cenozoic
Chiromyoides, as mentioned by Patterson, or the living Daiiben-
tonia, become even less possibly significant in the light of what is
now known of the skull. The skull is unknown in Chiromyoides,
but its upper incisors are entirely unlike those now known in Groe-
beria and the skulls of other plesiadapids are also quite dilTerent
(see especially Russell, 1964). There is an interesting resemblance
between the short, deep faces of Daubentonia and Groeberia, but
this is quite clearly functionally convergent and the two diff'er
markedly in other respects and in facial details. Almost all other
placental orders have basic diagnostic features absent in Groe-
beria or strongly contradicted in this genus.
If Groeberia was a placental, it must almost perforce have
evolved independently and uniquely from ancestors as primitive
and undifferentiated as those now known from the late Cretaceous
and, in decreasing numbers and generality, quite early Cenozoic.
Here one turns to the indirect and yet cogent evidence: that no
such placentals are known from South America; that equally pri-
mitive and undifferentiated marsupials are known from there; that
those marsupials did diverge in independent and unique lines; and
that derivation of Groeberia from a potent and definitely South
American source is at least a likely hypothesis.
Patterson's views on the affinities of Groeberia were buttressed
by evidence for referring it to the Caenolestoidea. That evidence.
1970 GROEBERIA 13
from the single poorly preserved fragment of mandible then known,
was as follows:
1 . One greatly enlarged lower incisor, with enamel on an-
terior face and alveolus (or "parte basal") parallel to
median line of symphysis (not to the horizontal ramus or
tooth row).
2. Strong salient coronoid process.
3. Molars with short talonids with posterior entoconid and
hypoconid, united by a transverse crest, a short crista
obliqiia, and a shallow basin.
4. Molar trigonids with one lingual (probable metaconid)
and two labial cusps (probable paraconid and protoco-
nid), as in caenolestoids except Caenolestinae.
5. Trigonids and talonids subequal, as in Palaeothentinae
and Abderitinae.
6. Masseteric crest absent; very poorly defined in Palaeothen-
tinae.
7. Inflection of lower border beginning anterior to cheek
teeth; usually posterior to cheek teeth in Caenolestoidea
but beneath M^ in Parabderites bicrispatus.
This was a valid analysis of the admittedly deficient evidence,
but the conclusion may be retroactively queried in the light of
present knowledge. As to (1 ), no unquestioned caenolestoids have
hypselcdont incisors comparable to these in form or function, but
the placing of the alveoli is an interesting point. (2) is not par-
ticularly diagnostic. Regarding (3)-(5), the molar structure is
difiicult to make out on the specimen, but I believe that Patterson
has correctly interpreted it. However, it diff'ers characteristically
from probably ancestral didelphid structure in litde more than the
more labial position of the paraconid, an occlusal adjustment that
could well arise more than once. (6) is somewhat dubious and
is not diagnostic for caenolestoids. (7) dift'ers from caenolestoids
more than it resembles them.
A reasonable but inconclusive case was presented on the basis
then available and pending acquisition of further knowledge. The
still quite limited further knowledge now acquired does not flatly
contradict that case, but neither does it strengthen it, and it even
weakens it to some extent. I see no special resemblance of known
parts of G. pattersoni to any unquestionable caenolestoid. On the
contrary, the extreme abbreviation of the face, the reduction of
the incisors to two, and their truly gliriform, hypselodont nature
14 BREVIORA No. 362
are almost the opposite of known trends in the Caenolestoidea.
Even the Polydolopidae, superficially most seemingly rodentlike
of undoubted caenolestoids, are not really very rodentlike in habi-
tus and evolved in a direction very unlike that of Groeberia (see
especially Simpson, 1948; Paula Couto, 1952).
Patterson (1952: 6) who com'pdL^td Groeberia \n\\\\ Argyrokiiius
and concluded that they are not specially related beyond their both
being marsupials. With greatly increased information on Argyro-
logiis, I agreed (Simpson, 1970), and the present addition to
knowledge of Groeberia does not change that opinion. Indeed, the
contrasts between Argyrolagiis and Groeberia, both somewhat
rodentlike, are remarkable. Argyrolagiis has an extremely elon-
gate, shallow rostrum and face, very posterior orbit, short, shal-
low zygoma, hypselodont cheek teeth. Groeberia has extremely
short, deep rostrum and face, very anterior orbit, long deep
zygoma, brachydont cheek teeth.
Almost complete knowledge of dentition and skeleton of Argyro-
lagiis contradicted previous opinion that it might be a caenolestoid
and required placing it in a separate superfamily. I suspect that
the same might happen if we had equally good information on
Groeberia, but we do not. This knowledge is still so scanty that
I believe it would be unreasonable or, at best, premature at this
point to classify the Groeberiidae other than as Marsupialia in-
certae sedis.
Proposals have long been made to divide the marsupials into
suborders, and recently, to divide them into two or more orders.
If suborders Polydactyla and Syndactyla were recognized, I would
predict that discovery of foot bones would aline Groeberia with
the polydactyls. If Polyprotodonta and Diprotodonta were recog-
nized, Groeberia would be descriptively, typologically, or pheneti-
cally diprotodont, but I believe that would be profoundly mis-
leading because it inevitably suggests connection with the Austral-
ian diprotodonts, and Groeberia is not so incertae sedis as all
that. It has no suggestion at all of Australian affinities. The
ordinal system of Ride ( 1964) has no sure ordinal place for it, as
the evidence that it may be a "paucituberculate" (caenolestoid)
is insufficient, but reference to the Marsupicarnivora would be
rather anomalous (although like all marsupials it doubtless arose
from the group so named by Ride), and pertinence to the Pera-
melina or Diprotodonta (seiisii Ride) is out of the question.
Kirsch's (1968) arrangement also has no sufficiently likely place
for Groeberia in any one of his orders.
1970 GROEBERIA 15
Bioloij;}'. In the absence of postcranial remains, nothing can
be safely inferred as to body build, hmb proportions, or locomo-
tion in Groeberia. Biological inferences are further limited by the
absence of specimens of the neurocranium and by the imperfec-
tions of the only two specimens known. The following are the
principal characters of probable functional importance that are
known:
1 . Face and snout short.
2. Face and mandible deep.
3. infraorbital foramen small.
4. Orbit relatively anterior.
5. Orbit of moderate size.
6. Masseteric origin on zygoma.
7. Heavy hypselodont incisors with labial enamel.
(S. 1' recumbent.
9. Short lower diastema near alveolar level.
10. Comparatively small, brachydont cheek teeth.
1 I . Large coronoid process.
12. Small (no?) masseteric crest.
13. Inverted angle.
The habitus is rodentlike at first sight, but no living rodent and
in fact no other animal, living or fossil, known to me combines
all those characteristics. 3, 6, 11, and 13 are usual and 4, 5, and
10 common in marsupials; all but 13 are also fairly common in
placentals. 1, 2, 7 and 8 occur in the other known marsupials
that are most rodentlike, the Australian wombats. They are lack-
ing in caenolestoids, also somewhat but much less rodentlike in
some genera. With only the partial exception of 13, all these
characters occur in one placental rodent or another, but not in
this combination. For example some caviomorphs, such as
Echbnys, have characters 5, 7, 8, 10 and a functional modification
of 13. but notably differ in 3, 6. 9. 11 and 12. Aplodontia has
3, 4, 6, 7, moderate 8, I 1 and moderate 12, but differs markedly
in 1,2, 5, 9, 10, and 13. Xenis (a sciurid) agrees well in 1,
3, 4, 7, 8, and 10, but less well in 2 and 9, and not at all in
5, 6, 11, 12, and 13.
There is no doubt that Groeberia was a powerful gnawer, at
least as much so as hares, rodents, wombats, or Daubentonia. Its
incisors were not merely a pincer apparatus as in all known caeno-
lestoids and all diprotodonts (phalangeroids) except the wombats.
They are decidedly more adapted to gnawing than in the argyro-
lagoids, even though the latter, unlike caenolestoids, also have
16 BREVIORA No. 362
hypselodont incisors. Gnawing is strongly connected with food
gathering in recent animals, but not exclusively so. On the other
hand, Groeheiias small area of brachydont cheek teeth is very
different from the hypselodont teeth of wombats, argyrolagoids,
and many rodents, and is more nearly comparable with the cheek
dentition of caenolestoids, squirrels, and Daubentonia. The com-
bination suggests a food obtained by gnawing but prepared for
deglutition by crushing or comminution without grinding. Pos-
sibilities are bark or wood-boring insects or fruits or nuts with
hard shells. However, some murids with strong gnawing apparatus
and limited, brachydont cheek teeth are virtually omnivorous or
even carnivorous. I see no way to correlate Groeberias unique
combination of characters with an equally unique diet or with
any closely specific way of life.
No rodents are known in the Divisadero Largo or any earlier
South American formation. Simpson, Minoprio, and Patterson
(1962: 289) mentioned the possibility that the presence of the
rodentlike marsupial Groeberia indicated age before rodents
reached that area, hence pre-Deseadan. The ecological aspect of
that suggestion would still hold even if the determination of the
relative time of entry of rodents proved to be incorrect. (There
is still no opposing evidence.) The rodent habitus surely evolved
elsewhere and was in being when the first rodents entered South
America, whether in the Deseadan or, as is quite likely, somewhat
earlier. On the other hand, the ancestral habitus of Groeberia
almost certainly evolved in South America before rodents reached
there, convergent to some extent toward the absent rodents and
entering niches with some points of similarity.
Zoogeography. It is the most reasonable hypothesis that the
Groeberidae did evolve in South America, even though their pos-
sible relationship to the Caenolestoidea is quite dubious, and there
is no suggestion of special descendant relationship (involving
specialization) to any other South American group. Origin from
South American didelphoids is as likely as any other, if not more
so. There is no special resemblance to any Australian marsupials
suggestive of genetic affinity beyond the remote ancestry of all
marsupials. No marsupials are known from Africa and any idea
of connection there would be purely gratuitous at present.
Nevertheless it is strange that three of the most peculiar, most
specialized known groups of South American marsupials appear
in the presently available record without known ancestors, only to
1970 GROEBERIA 17
vanish again immediately (geologically speaking) or soon there-
after: Groeberiidae, only in the Divisadero Largo formation; Nec-
rolestidae, only in the Santa Cruz formation; and Argyrolagidae,
only from Huayquerian to Uquian. A possible clue is that all
these faunas are in Temperate Zone Argentina and that earlier
faunas are as yet very inadequately known farther north on the
continent. It is a reasonable hypothesis, as yet without direct
evidence, that these groups evolved in what are now (and quite
likely were then) the tropics and are picked up in our record only
when they spread rather briefly to what was for them a marginal
area.
REFERENCES
KiRSCH, J. A. W. 1968. Prodromus of the comparative serology of
Marsupialia. Nature, 217: 418-420.
Pascual, R.. E. J. Ortega Hinojosa, D. Gondar, and E. Tonni. 1965.
Las Edades del Cenozoico mamalifero de la Argentina, con especial
atencion a aquellas del Territorio Bonaerense. Prov. de Buenos Aires,
Sep. An. Com. Invest. Cient. Bs. As., VI: 165-193.
Patterson. B. 1952. Un nuevo y extraordinario marsupial deseadiano.
Rev. Mus. Municipal Cien. Nat. y Tradic, Mar del Plata, 1: 49-44.
Paula Couto, C. de. 1952. Fossil mammals from the beginning of
the Cenozoic in Brazil. Marsupialia: Polydolopidae and Borhyaenidae.
American Mus. Novitates. No. 1559: 1-27.
Peyer, B. 1968. Comparative Odontology. (R. Zangerl. ed.) Chicago,
Univ. of Chicago Press.
Ride. W. D. L. 1964. A review of Australian fossil marsupials. Jour.
Roy. Soc. Western Australia, 47: 97-131.
Russell, D. E. 1964. Les Mammiferes Paleocenes d'Europe. Memoires
du Mus. Nat. D'Histoire Nat.. Serie C, Sciences de la Terre, XIII:
1-324.
Simpson, G. G. 1948. The beginning of the Age of Mammals in South
America. Bull. American Mus. Nat. Hist., 91: 1-232.
, 1970. The Argyrolagidae. extinct South American
marsupials. Mus. Comp. Zool.. 139 (1): 1-86.
Simpson. G. G., J. L. Minoprio, and B. Patterson. 1962. The mam-
malian fauna of the Divisadero Largo formation, Mendoza, Argentina.
Bull. Mus. Comp. Zool.. 127 (4): 239-293.
BREVIORA
Miiseiuiiti of Compsirative Zoology
Cambridge, Mass. 8 January, 1971 Number 363
NON-SPECIFICITY OF HOST-SELECTION IN THE
ECTOPARASITIC SNAIL ODOSTOMIA (MENESTHO)
BISUTURALIS (SAY) (GASTROPODA: PYRAMIDELLIDAE)
Robert C. Bullock and Kenneth J. Boss
Abstract. Ectoparasitic pyramidellid gastropods have often been
considered host-specific, although a few species have been reported to feed
on a variety of hosts under laboratory conditions. A large population of
Odostomia (Menestho) bisuturalis (Say) at Duxbury Beach, Massachu-
setts, provided an ideal opportunity to study the association of this ecto-
parasite with the various moUusks found in the vicinity. Our results re-
vealed that O. bisuturalis was associated with seven species of moUusks,
most commonly Nassariiis obsoletiis (Say) and Mytilus editlis Linnaeus.
The great abundance of O. bisuturalis and the fact that Crassostrea vir-
ginica (Gmelin), its usual natural host, was absent from the study area,
appear to account for the non-host-specificity observed.
INTRODUCTION
The host-specificity of the ectoparasitic snails of the family
Pyramidellidae has been the subject of several papers during the
last decade. Early observations led some workers, notably Fretter
and Graham (1949; 1962), to state that these ectoparasites were
host-specific. More recent studies have revealed that under labora-
tory conditions certain pyramidellids actually feed on a variety of
hosts (Ankel and Christensen, 1963; Scheltema, 1965; Robertson,
personal communication), although possible host-preference may
be a factor (Boss and Merrill, 1965). The observations reported
here suggest that some species of pyramidellids are not host-
specific.
What actually constitutes a true parasitic relationship has been
questioned (Robertson and Orr, 1961; Dehlinger, unpublished
MS). While observations of pyramidellids associated with
2 BREViORA No. 363
various organisms indicate, by their proximity or physical contact,
possible parasite-host relationships, most authors have been
wary of such evidence. Robertson (personal communication)
stated that the presence of an Odostomia on a possible host may
only reflect the pyramidelUd's need for a suitable substrate. This
view is supported by some of our observations and those of Schel-
tema (1965), who noted O. bisuturalis clinging to the under sur-
faces of stones in a region where Littorina littorea L., a "labora-
tory host" for this species, was abundant. Recent workers have
indicated what constitutes a true parasitic relationship: insertion
of the proboscis and action of the buccal pump (Robertson, per-
sonal communication; Scheltema, 1965) or when the parasite is
"less than V& in. from the edge of the mantle of the host"
(Boss and Merrill, 1965). Although the former is a more exact
method, we have necessarily followed Boss and Merrill (1965),
since our observations were made during the low tide period when
the ectoparasites were not actively feeding on exposed hosts.
OBSERVATIONS
An abundance of Odostomia (Menestho) bisuturalis (Say,
1822) was noted on the tidal flat on the harbor side of Duxbury
Beach, Massachusetts, in June, 1969. Sampling at several stations
along the beach during a -1.6 tide on 3 June and subsequent
visits to the study area provided ample field evidence of the occur-
rence of this snail on numerous intertidal moUuscan hosts.
The Odostomia were not limited to a particular region of the
tidal flat, for they were found from below the low-water mark to
the upper portion of the intertidal zone. Quantitative samples
were taken in the region at the east end of the Duxbury Beach
bridge. All moUusks collected were carefully examined for the
presence of Odostomia. O. bisuturalis, the only pyramidellid
found, was collected from seven different species of moUusks:
Littorina littorea (L.), Urosalpinx cinereus (Say), Crepidula con-
vexa Say, Crepidula fornicata (L.), Nassarius obsoletus (Say),
Nassarius trivittatus (Say), and Mytilus edulis Linnaeus. They
were also observed on the egg capsules of Nassarius sp. and Poli-
nices sp., and on empty shefls.
O. bisuturalis was most abundant on three hosts: Mytilus edulis,
which formed large mats on the mud flats, and on the snails Nas-
sarius obsoletus and Littorina littorea, which occasionally had
1970 ECTOPARASITIC ODOSTOMIA 3
from one to three ectoparasites on the operculum or on the Hp of
the shell. The Mytilus that were observed in water often had a
number of ectoparasites situated on the margins of the mantle,
away from the hinge, similar to the position assumed in Odostomia
scalaris MacGillivray on Mytilus in Europe. However, most of
the Mytilus population at Duxbury was exposed at low tide and
the ectoparasites had migrated to moist areas within the Mytilus
mat. In the laboratory we have observed O. bisuturalis feeding
on Mytilus.
Table I shows the relative abundance of Odostomia on three
hosts. In the case of Nassarius obsoletus several items should be
noted: 1) the parasites occurred on samples with a large mean
length in a frequency of one Odostomia to three or four Nassarius;
2) smaller individuals of Nassarius were significantly less para-
sitized, with only one parasite per ten individuals. This latter ob-
servation was also noted in studies of O. impressa (Say) (Hop-
kins, 1956) and O. dianthophila Wells and Wells (Wells and
Wells, 1961). At one station A', obsoletus was present in large
numbers, possibly 12,000/m-, with an equally high concentration
of O. bisuturalis, 7,000/m-. These figures indicate that at times
O. bisuturalis, a usually overlooked organism, must play a mod-
erately significant role in the flow of energy in a tidal flat com-
munity. In other species, not quantitatively sampled, the occur-
rence of Odostomia was even greater, in some cases amounting
to two ectoparasites per host, e.g., Littorina.
Previous published records of field observations have not re-
vealed O. bisuturalis in association with any of the seven species
of moUusks reported in this study. In the laboratory, O. bisutu-
ralis is known to feed on: Littorina littorea (L.) (Scheltema,
1965; Robertson, 1967); Bittium altematum (Say) (Scheltema,
1965); Crucibulum striatum (Say) and Crepidula fornicata (L.)
(Boss and Merrill, 1965). Previously, the only known natural
host of O. bisuturalis was the American oyster, Crassostrea vir-
ginica (Gmelin) (Loosanoff, 1956; Boss and Merrifl, 1965), a
species not present at Duxbury Beach.
DISCUSSION
These observations demonstrate that at least one pyramideflid
is not host-specific. Further, laboratory studies have shown that
certain Odostomia may have numerous hosts. Thus, should these
4 BREVIORA No. 363
species actually be non-host-specific, as O. bisiituralis seems to
be, an examination of all environmental factors involved in host-
specificity and host-preference becomes imperative.
The great abundance of O. bisiituralis led to the consideration
of the question of density-dependent factors in feeding behaviour.
While our observations show the non-specific host-selection of
O. bisuturalis, they do not reveal any specific information con-
cerning host-preference. An abnormally large population of O.
bisuturalis or a diminished preferred food source might force
many individuals to feed on hosts they would otherwise reject.
The biological relationship between a motile ectoparasite and
host-species can be likened to that between a predator and its
prey. The odostomia-type of predator-prey relationship with re-
spect to food preference is commonly observed in animals not
totally dependent on one organism for survival. It has been
demonstrated that total food abundance, relative abundance of
food types, spatial distribution of foods and predator satiation
afl'ect feeding preferences in some animals (Ivlev, 1961). In mol-
lusks, for example. Wells (1958) found that although the oppor-
tunistic gastropod Fasciolaria hunter ia (Perry) prefers to eat the
small oyster drill, Urosalpinx, it will eat oysters if only a minimal
number of the more desirable gastropods are present. Further, in
a study of the relationship between time and energy in food pref-
erences, Emlen (1966: 617) suggested that: 1) ''Animals should
be more selective in their choice of foods when satiated or when
food is common, more indiscriminate when starved or when food
is scarce"; and, 2) "Food preferences appear to change readily
and appropriately to changes in the environment."
The particular circumstances in which the ectoparasitic popula-
tion existed were unusual in that: 1 ) there was a very large popula-
tion of ectoparasites; and, 2) the preferred natural host for this
species, Crassostrea virginica, was not present in the local eco-
system. Thus, our observations corroborate the hypotheses of
Emlen and indicate that Odostomia bisuturalis may have various
host-species under natural conditions.
1970 ECTOPARASITIC ODOSTOMIA 5
ACKNOWLEDGMENTS
The manuscript was critically read by Messrs. R. I. Johnson
and M. K. Jacobson and Dr. R. D. Turner. Mr. W. Baranowski
brought our attention to the occurrence of great numbers of Odo-
stomia at the Duxbury site, and Mr. S. Britz measured the speci-
mens.
BREVIORA
No. 363
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1970 ECTOPARASITIC ODOSTOMIA 7
REFERENCES CITED
Ankel, F., and a. M. Christensen. 1963. Non-specificity in host selec-
tion by Odostomia scalaris MacGillivray. Vidensk. Medd. fra Dansk
naturh. Foren., 125: 321-325.
Boss, K., AND A. Merrill. 1965. Degree of host specificity in two species
of Odostomia (Pyramidellidae: Gastropoda). Proc. Malac. Soc. Lon-
don, 36: 349-355.
Dehlinger, R. 1969. Host specificity and the feeding ecology of the
ectoparasitic family Pyramidellidae. Unpublished manuscript, 55 pp.
Emlen, J. M. 1966. The role of time and energy in food preference.
Amer. Nat., 100: 611-617.
Fretter, v., and a. Graham. 1949. Feeding and reproduction in pyra-
midellids. Nature, 163: 361-362.
. 1962. British Prosobranch Molluscs. London, Ray
Society, 755 pp.
Hopkins, S. H. 1956. Odostomia impressa parasitizing southern oysters.
Science, 124: 628-629.
Ivlev, V. S. 1961. Experimental Ecology of the Feeding of Fishes. New
Haven, Yale Univ. Press, 302 pp.
Loosanoff, V. L. 1956. Two obscure oyster enemies in New England
waters. Science, 123: 1119-1120.
Robertson, R. 1967. The life history of Odostomia bisiitiualis, and
Odostomia spermatophores (Gastropoda: Pyramidellidae). Year Book
Amer. Phil. Soc, 1966: 368-370.
Robertson, R., and V. Orr. 1961. Review of pyramidellid hosts, with
notes on an Odostomia parasitic on a chiton. Nautilus, 74: 85-91.
Scheltema, a. H. 1965. Two gastropod hosts of the pyramidellid gas-
tropod Odostomia bisiitiiralis. Nautilus, 79: 7-10.
Wells, H. W. 1958. Predation of pelecypods and gastropods by Fas-
ciolaria hunteria (Perry). Bull. Mar. Sci. Gulf and Caribbean, 8:
152-166.
Wells, H. W., and M. J. Wells. 1961. Three species of Odostomia
from North Carolina, with description of new species. Nautilus, 74:
149-157.
BREVIORA
MiaseitairM of Comparative Zoology
Cambridge, Mass. 8 January, 1971 Number 364
A NEW SCINCID LIZARD FROM BOUGAINVILLE,
SOLOMON ISLANDS
Allen E. Greer and Fred Parker^
Abstract. The relationships of Sphenomorphus transversus, n. sp.,
from Bougainville, Solomon Islands, are obscure, but in squamation it is
most similar to inaciilatiis, boiilengcri, forinosensis, Uneopiinctiilatiis. and
indiciis from eastern Asia; nielanochlorus from New Guinea; and sanctiis
from Sumatra and Java. S. transversus differs most noticeably from these
species and from other Bougainville skinks in its dorsal pattern of trans-
verse dark brown bands on a light olive ground color.
During investigations in 1960-1963 by Parker on Bougain-
ville, Solomon Islands, a single individual of a previously unde-
scribed species of skink was collected. Since one subsequent trip
(1966) has failed to uncover other specimens of the species, and
as the possibilities of a second return trip to Bougainville in the
near future are slim, it seems best to describe the new species from
the single specimen at hand.
On the basis of current generic concepts, the species is assigned
to the genus Sphenomorphus and may be known as
Sphenomorphus transversus^ new species
Holotype. Museum of Comparative Zoology 76485; collected
by a native for Fred Parker at about 2000 feet above sea level in
an area approximately five miles east of Kunua, northeastern
Bougainville (Fig. 1), on 9 September 1962.
1 P. O. Box 52, Daru, Western District, Territory of Papua and New
Guinea.
- The species name calls attention to the dark transverse bars on the
dorsum.
BREVIORA
No. 364
155° E
56° E
BUKA I
30 miles
■6°S
•7°S
6°S-^
I55°E
SHORTLAND
MONO I (^
FAURO
7°S-
156° E
Figure 1. Map of Bougainville showing the location approximately 5
miles east of Kunua where the type and only known specimen of Spheno-
morphiis transversus was collected.
Diagnosis. Similar in squamation to those skinks of the genus
Sphenomorphus (Table 1 ) that have a single anterior loreal, the
frontal in contact with 3 or more of the 5 or more supraoculars,
frontoparietals and interparietal distinct, no nuchals or trans-
versely enlarged scales in the two vertebral rows (Figs. 2 and 3),
I97'
A NEW SOLOMON ISLANDS SKINK
and the digits and limbs well developed and overlapping when
adpressed to the body, but differing from other skinks with this
diagnosis in having the following combination of characters: pre-
frontals separated medially, 36 rows of smooth scales around mid-
body, 28-29 smooth subdigital lamellae on the 4th toe, and a
color pattern of brown transverse bands on a light olive-green
ground color (Figs. 4 and 5) — a color pattern most similar to
those of the distantly related Sphenomorphus flavipes, Scincella
prehensicaiida, and Leiolopisma semoni of New Guinea.
Description. Body form relatively slender; well-developed pen-
tadactyl digits and Umbs that overlap when adpressed to the body
(tip of 4th toe reaches middle of forearm); snout-vent length 68
mm, tail 92 mm.
Figure 2. Dorsal view of the head of the holotype of Sphenomorphus
transversus (MCZ 76485).
BREVIORA
No. 364
Head not depressed, snout somewhat pointed; rostral slightly
wider than deep, projecting slightly onto dorsal surface of snout
between nasals; external naris in single nasal; no supranasals;
single anterior and posterior loreals; frontonasal slightly wider
than long, forming a short suture with the rostral and a very
short suture with the frontal; prefrontals large, barely separated at
their inner angles; frontal 1% times as long as wide, in contact
with the three anteriormost supraoculars; 5 supraoculars, the
first smallest, but in no way to be confused with the anterior super-
ciliaries; lower eyelid scaly; 6th supralabial most directly below
eye; a complete row of subocular scales separates scales of lower
eyelid from supralabial series; frontoparietals paired and subequal
in size with the single interparietal that is sharply pointed pos-
teriorly; parietals meeting behind interparietal and bordered
posteriorly by a single large temporal on either side and 5 large
dorsal scales between the temporals; no symmetrical series of
nuchals.
External ear opening vertically elliptic, without auricular lobes;
tympanum sunk slightly below level of skin; 36 smooth scales
around midbody, the scales of the two vertebral rows not larger
than those of the immediately adjacent rows; a pair of enlarged
preanals; scales of three median subcaudal rows subequal in size.
Digits rather long and slender; subdigital lamellae smooth and
undilated throughout length of digit; 28-29 lamellae beneath 4th
(longest) toe; upper surface of 4th toe covered by one or two
Figure 3. Lateral view of head of Sphenoworpliiis tnnisversus (holo-
type).
1971 A NEW SOLOMON ISLANDS SKINK 5
single scales at distal end, 3 rows of scales throughout center part
and 4 rows near base (see Brongersma, 1942).
Dorsal ground color light olive with a series of complete and
incomplete transverse dark brown bands from nape to base of
tail, the brown bands terminating in slightly expanded blotches
on sides (Fig. 5), an effect especially pronounced at midbody; a
horizontal brown stripe from anterior loreal through eye to tem-
poral region; brown blotches on anterior and upper surfaces of
limbs as well as on upper surface of tail; venter immaculate except
for a few faint brown spots on throat and underside of tail.
In life the undersides of the limbs, body, and tail were bright
yellow.
Field Notes. The only known specimen of S. transversus was
taken by a native collector under a decaying log on the steep side
of a montane river valley covered with tall primary forest. The
natives did not recognize it as being distinct from S. concinnatus,
a species common at the type locality of 5. transversus.
Morphological Comparisons with Other Bougainville Skinks.
S. transversus is immediately distinguishable from S. concinnatus
by its more sharply tapered, longer snout; the absence of a dark
blotch between the ear opening and the forellmb; the smaller ex-
ternal ear opening; and, the regular transverse barring.
Only two other Bougainville skinks, S. taylori and S. cranei —
both very different from S. transversus in squamation — have
transverse bands on the dorsum. In both these species, however,
the dorsal pattern consists of very light transverse bands on a dark
ground color, whereas in S. transversus the transverse bands are
darker than the ground color. S. transversus also has a more
noticeably pointed snout than either S. taylori or S. cranei.
Skull Characters. It is extremely difficult, if not impossible,
to remove the skull of most skinks without severely damaging the
skin of the head. For this reason we have not attempted to pre-
pare a skull from the type and only known specimen of S. trans-
versus. We have, however, had a palatal view of the skull, and
the salient features are as follows: there are 9 premaxillary teeth;
the palatine and pterygoid bones meet along the midhne to form
a fairly extensive secondary palate; there is no ectopterygoid pro-
cess; and, there are no pterygoid teeth. Unfortunately, these
characteristics are not particularly diagnostic, for they would not
exclude S. transversus from close relationship with any number
6 BREVIORA No. 364
of other lygosomines, including those discussed below that are
most like S. transversiis on the basis of external morphology.
Comparison with Morphologically Similar Species. In squama-
tion S. transversus is most similar to those species of Sphenomor-
phus (Table 1) that have a single anterior loreal, the frontal in
contact with 3 or more of the 5 or more supraoculars, the fronto-
parietals and interparietal distinct, no nuchals or transversely en-
larged scales in the two vertebral rows, and the digits and limbs
well developed and generally overlapping when adpressed to the
body. This assemblage, which is almost surely not monophyletic,
is distributed from southern Asia through the Indo-Australian
archipelago and Philippines to New Guinea, but not Australia.
Seven species in this group have the prefrontals separated me-
dially (in all or some individuals), as is the case in the single
specimen of S. transversus. Six of these seven species (maculatus,
boulengeri, jormosensis, lineopimctulatus , and indiciis from
southern Asia and melanochlorus from New Guinea) have sub-
stantially fewer subdigital lamellae on the 4th toe (16-22) than
does transversus (28-29), and the seventh {sanctus from Suma-
tra and Java) has finely striated body scales to distinguish it from
the smooth-scaled transversus. Furthermore, none of these seven
species have a dorsal body pattern consisting of well-defined dark
crossbars as does transversus.
This very characteristic dorsal body pattern of dark crossbars
on a light ground color is most similar to the patterns of the cross-
banded color morph of Sphenomorphus fiavipes, the females of
Scincella prehensicauda, and of all Leiolopisma semoni. These
three species are endemic to neighboring New Guinea and might,
therefore, seem to be likely relatives of Sphenomorphus trans-
versus. Current work on scale and palatal characters, however,
indicates that while fiavipes, prehensicauda and semoni are them-
selves closely related (in spite of current generic allocations), they
are only distantly related to SphenomorpJius transversus.
ACKNOWLEDGEMENTS
Dr. Ernest E. WiUiams of the Museum of Comparative Zoology
read the manuscript in several drafts and offered several helpful
suggestions. Mr. Laszlo Meszoly did the drawings for Figures 2
and 3, and Mr. Tan T. Riddell took the photographs for Figures
1971 A NEW SOLOMON ISLANDS SKINK 7
4 and 5. Part of Greer's work on this paper was done while he
was a postdoctoral fellow of the National Science Foundation.
Partial support was provided by National Science Foundation
grant GB 6944 to Ernest E. Williams.
LITERATURE CITED
Brongersma, L. D. 1942. On the arrangement of the scales on the
dorsal surface of the digits in Lygosoma and allied genera. Zoologische
Mededeelingen. 24 (1-2): 153-158.
BREVIORA
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BREVIORA
No. 364
Figure 4. Dorsal view of the holotype of Splicnomorphiis transvcrsus
(MCZ 76485) showing the dark transverse bands which give the species
its name.
1971
A NEW SOLOMON ISLANDS SKINK
11
Figure 5. Lateral view of Sphenomorphiis transversiis (holotype).
BREVIORA
Me seem of Comparative Zoology
Cambridge, Mass. 15 January, 1971 Number 365
CHARACTERS AND SYNONYMIES AMONG THE GENERA
OF ANTS. PART IV. SOME GENERA OF SUBFAMILY
MYRMICINAE (HYMENOPTERA: FORMICIDAE)
William L. Brown, Jr.^
Abstract. Archaeomyrmex is a new junior synonym of Myrmecina,
and tribe Archaeomyrmicini accordingly is a synonym of Myrmecinini.
Dodoiis is a new junior synonym of Pristomyrmex, and the synonymy of
Hylidris under Pristomyrmex is reaffirmed. Limnomyrmex is a new junior
synonym of Leptothorax subgenus Nesomyrmex.
The genera considered in this part all belong to subfamily Myr-
micinae. My main purpose here is to explain briefly why some
new synonymy should be proposed. The taxa concerned have all
been studied during the course of the project "a reclassification of
the Formicidae," supported by National Science Foundation Grants
G-23680, GB-2175, and GB-5574. The taxonomic conclusions
will eventually be embodied in a synopsis and illustrated keys to
the ant genera of the world. I feel that justification for revisionary
changes should be published as the need for the changes becomes
clear. In this way, important findings of the study are made avail-
able for use by all ant taxonomists without undue delay, and the
synoptic parts can be freed from the clutter of many detailed
taxonomic arguments.
Myrmecina
Myrmecina Curtis, 1829, Brit. Entom. 6: 226, pi. 265, male. Type
species by original designation Myrmecina latreillii := Formica graminicola.
Arclweomyrmex Mann, 1921: 448-451. Type species: Archaeomyrmex
cacabau, by original designation. NEW SYNONYMY.
1 Department of Entomolo{!;y, Cornell l'ni\ ersity, Ithaca, New York, 14850.
2 BREVIORA No. 365
The unique type of A. cacabau has been searched for in vain in
the U. S. National Museum and Museum of Comparative Zoology
ant collections, and must be considered lost. Fortunately, Mann's
description and figures are reasonably detailed. From them, it is
clear that the species is essentially a Myrmecina, a fact acknowl-
edged by Mann when he wrote: "The epinotal and petiolar struc-
ture are not unlike certain species of Myrmecina." Mann empha-
sized the ventrolateral carina on each side of the head, probably
without realizing that this is an invariable character of Myrmecina.
He also cited the partly smooth, partly costate sculpture of the
body, especially the trunk, which does set this species off from the
"average" Myrmecina. But Myrmecina forms with more or less of
the head and trunk smooth have not been completely unknown,
and they even occur as phenetic variants in species that are usually
heavily sculptured in these regions.
Several Myrmecina species (at least three of them still unde-
scribed) with predominantly smooth and shining head and trunk
are in the MCZ collection. Two of the undescribed species are
from the New Hebrides; in each of these, the posterior pair of
propodeal teeth is spiniform, but the coarse costate sculpture of
the lower pleural areas is preserved, as it is in cacabau. Another
smooth species has come to me from the Philippines. The geo-
graphical and morphological gaps are thus closed between the
Fijian and Indo-Australian representatives of what is obviously a
single stock. A complete examination of the A. cacabau descrip-
tion fails to reveal any character that will separate it from Myrme-
cina at generic level. Together with the two New Hebrides species,
cacabau should be taken to represent no more than a weak species-
group of Myrmecina.
Incidentally, an interesting characteristic of some of these species
is the extreme reduction of antennomere III, just distal to the
pedicel. The distinction between " 1 1 -segmented" and "12-seg-
mented" antennae in these forms may prove impossible to maintain
as a key character.
Mann's tribe Archaeomyrmecini of course falls as a new
synonym of Myrmecinini.
Pristomyrmex
Pristomyrmex Mayr, 1866: 903. Type species: Pristomyrmex piingens,
monobasic.
1971 SOME GENERA OF MYRMICINE ANTS 3
Hylidris Weber, 1941: 190. Type species: Hylidris myersi, by original
designation. — 1952: 15-22. Synonymized by Brown, 1953: 9-10.
Dodous Donisthorpe, 1946: 145; worker, male, larva. Type species:
Dodous trispinosus, by original designation. NEW SYNONYMY.
Dodous was based on the single species trispinosus, from Cocotte
Mountain, Mauritius. Syntypes of this species deposited in the
British Museum and in the Museum of Comparative Zoology are
basically Pristomyrmex in form, but they have an extra pair of
small teeth on the mesonotum, and the head and trunk are finely
and regularly costulate (= striate). The larva, rather vaguely
figured in the original description (fig. 4), has the elongate, pro-
tuberant head characteristic of Myrmecina and Pristomyrmex.
Donisthorpe also figured the male, which is like known Pristomyr-
mex males in general habitus. The genitalia as very briefly de-
scribed may be aberrant, but then the genitalia have not been
studied at all in most other Pristomyrmex species.
A second species of Dodous, D. bispinosus, was described from
Mauritius by Donisthorp)e in 1949. I collected a small sample of
strays of this species at the type locality, Le Pouce (Mountain) on
1 April 1969, the last day of a short stay on Mauritius. The speci-
mens were all foraging workers taken on trees and on the main
path through the scrubby native forest at about 800 m elevation
on the "plateau^' near the summit. (A sudden storm prevented my
finding any nests.) D. bispinosus differs from D. trispinosus in
lacking the mesonotal teeth (though actually some of my bispinosus
specimens have low mesonotal tubercles in place of the teeth) and
in having predominantly smooth and shining sculpture. In fact,
D. bispinosus is a rather ordinary, if slightly long-legged, Pristo-
myrmex, and D. trispinosus goes only one step beyond. It seems
absurd to put these two closely related species in different genera.
Discovery of the annectant D. bispinosus makes it clear to me that
Dodous is only the Mauritian complement of the widespread Old
World genus Pristomyrmex. The concept of Dodous as a separate
genus is, as far as 1 am concerned, as dead as its namesake.
I have already (Brown, loc. cit.) placed Weber's Hylidris as a
synonym of Pristomyrmex. Weber opposed this synonymy, main-
taining that Hylidris is a distinct genus. But when he described
Hylidris, Weber took no note of the African species of Pristomyrmex
described previously by Santschi, Arnold, and Karavaiev, at least
some of which are senior synonyms of his own Hylidris species and
4 BREVIORA No. 365
subspecies (Weber, 1952). Weber has never produced a charac-
terization of Hylidris that will separate it as a genus from Pristo-
myrmex, and particularly from the long-synonymized "subgenus"
Odontomyrmex. \ have collected P. orbiceps in the Ivory Coast,
and can affirm that colony behavior (lethisimulation), larval form,
and general adult morphology are fully those of Asian and Aus-
tralian Pristomyrmex as I have seen them in nature.
Pristomyrmex is a sharply defined and compact genus, and there
is no reason that I know of to set the African species apart from it.
In fact, the African species are as nearly "average" for the genus
Pristomyrmex as one is likely to find.
Some other misconceptions expUcit or implied in Weber's dis-
cussion of 1952 need correction. The petiole of Pristomyrmex has
a distinct anterior peduncle, though it is short in some species. In
contrast, the related genus Myrmecina has a sessile petiole of more
or less prismatic shape. Pristomyrmex and Myrmecina are not
very closely related to tribe Tetramoriini, though two species have
been wrongly placed in tetramoriine genera in the past. The larvae,
for one thing, are very different, and it seems that they may furnish
the best tribal character for the Myrmecinini if we limit the tribe
to Pristomyrmex, Myrmecina, Acanthomyrmex, and possibly the
little-known Perissomyrmex of Guatemala, the larvae of which have
not yet been found. Tribe Tetramoriini is not "worldwide" in dis-
tribution, if one ignores obvious introductions by man. The tribe
has no native species in South or Central America, and only a single
species of Xiphomyrmex occurs in (Sonoran) North America.
Leptothorax subgenus Nesomyrmex
Nesomyrmex Wheeler, 1910, Bull. Amer. Mus. Nat. Hist., 28: 259.
Type species Nesomyrmex clavipilis, monobasic.
Leptothorax (Goniothorax) aiict., preoccupied.
Leptothorax (Caulomyrma) Forel, 1914, Bull. Soc. Vaudoise Sci. Nat.,
50: 233.
Limnomyrmex Arnold, 1948, Occas. Pap. Nat. Mus. S. Rhodesia, 2(14):
222. — 1952, Ibid., 2(17): 460, discussion. Type species Limiioiuyrmex
stramineiis, monobasic. NEW SYNONYMY.
Soon after its description, the late Dr. Arnold and I engaged in
correspondence on the question of the distinctness of Limnomyr-
mex from the subgenus Nesomyrmex of Leptothorax. He stoutly
maintained that Limnomyrmex was a good genus, and in 1952 {loc.
cit.) he argued again in print for this stand. Now that I have finally
1971 SOME GENERA OF MYRMICINE ANTS 5
seen the unique worker type of L. stramineiis in the Arnold Collec-
tion at Bulawayo, I can only place this species among the other
known African Nesomyrmex. I have searched in vain for any
characters that might set Limuomyrmex apart as a genus. In the
form of the trunk and both petiolar and postpetiolar nodes, it is
about "average" for a Nesomyrmex from Africa, and resembles in
a subdued way some neotropical members of the group. Sculptural
and a few other differences mark stramineus , but these do not seem
to be more than species characters. The antennae of stramineus
are 12-segmented; Nesomyrmex can have either 11 or 12 segments.
The status of Nesomyrmex as a subgenus of Leptothorax is main-
tained for the time being, pending the proper study of both taxa.
REFERENCES
Brown, W. L.. Jr. 1953. Characters and synonymies among the genera
of ants. Parti. Breviora No. 11: 1-13.
DoNisTHORPE, H. St. J. K. 1946. A new genus and species of Formicidae
(Hym.) from Mauritius. Proc. Roy. Entomol. Soc. London, ser. B,
15: 145-147.
Mann, W. M. 1921. The ants of the Fiji Islands. Bull. Mus. Comp.
Zool., 64: 401-499.
Mayr, G. 1866. Diagnosen neuer und wenig gekannter Formiciden.
Verhandl. Zool.-bot. Ges. Wien, 16: 885-908, pi. 20.
Weber, N. A. 1941. Four new genera of Ethiopian and Neotropical
Formicidae. Ann. Entomol. Soc. Amer., 34: 183-194.
. 1952. Studies on African Myrmicinae, I (Hymenoptera,
Formicidae). Amer. Mus. Novitates 1548: 1-32.
JBREVIORA
Mmseuinni of Comparsitive Zoology
Cambridge. Mass. 15 January, 1971 Number 366
PULSED SOUNDS OF THE PORPOISE
LAGENORHYNCHUS AUSTRALIS
William E. Schevil! and William A. Watkins^
Abstract. Pulsed sounds of the porpoise Lagenorhynchns aiistmlis of
southern Chile were recorded and analyzed. Most were low-frequency
clicks; some had a 2-kHz bandwidth centered near 1 kHz, and others had
a 10- or 12-kHz bandwidth with the principal frequency in the lower 5 kHz.
These porpoises also produced a rapidly pulsed tonal sound. All these
sounds were very low-level and rarely audible at a distance as great as 20 m.
We heard none of the whistlelike squeals characteristic of many delphinids.
From 12 November to II December 1968 the research ship
HERO of the National Science Foundation (Antarctic Research
Program) cruised between Valparaiso and Cape Horn, searching
for cetaceans and pinnipeds, mostly in the sheltered inland water-
ways of southern Chile. We were concerned with listening for and
recording the underwater sounds of these animals.
The species most frequently seen and collected was Lcigeno-
rhynchus australis (Peale, 1848 ), which has been reported from the
west coast of Chile south of about S Lat. 40° around Cape Horn
to the Falkland Islands. We heard them much less often than we
saw them.
Methods. The recordings that are analyzed here were made on
23 November in Canal Messier (at 48° 10' S) and 1, 3, 5, and 6
December west and south of Navarino Island behind Cape Horn.
An Atlantic Research LC-34 hydrophone was used to pick up the
sounds. An impedance-matching pre-amplifier (WHOI) was in-
serted in the cable 30 cm from the hydrophone. Two cable lengths
1 Contribution No. 2562 from the Woods Hole Oceanographic Institution.
2 BREVIORA No. 366
were used, 125 m from R/V HERO and 30 m from HERO's whale-
boat. The hydrophone depth varied with local conditions and ex-
periments, from 2 m to nearly 125 m; it was usually suspended 6-8
m from a surface float (a rubber balloon) and allowed to drift as
far from ship or boat as cable-length permitted.
Tape recordings were made with either a modified Uher 4400
recorder or a WHOI-built springwound recorder, using a hydro-
phone amplifier (Watkins, 1963). When the Uher was in use, the
system-response was limited to a bandwidth, within 4 db, of 40 to
20,000 Hz; with the WHOI machine, system-response was 20 to
32,000 Hz (within Vz db from 30 to 30,000 Hz). Playback for
analysis was on Crown 800 tape recorders. Spectrographic analy-
ses were made on a Kay Electric model 7029A analyzer and ampli-
tude analyses on a Tektronix 5 35 A oscilloscope.
The porpoises were approached as closely and as unobtrusively
as possible, but even so, often the only sounds heard from the ani-
mals were within the first 5 seconds of the listening attempts. Un-
fortunately, because of the disturbance of the water by arrival of
the boat and the motion of the hydrophone, a longer time than
this usually was required before local ambient noise could settle
down enough for faint sounds to be recorded. Usually nothing was
heard from the porpoises, partly because they were generally taci-
turn and seemed to produce sounds only occasionally, and partly
because their sounds were too faint to be audible except on close
approach, within a few meters of the animals. They appeared to
be silent when disturbed.
Sounds. The sounds heard from Lagenorhynchus aiistralis were
all pulsed. Mostly they were clicks produced in short series or slow
bursts, but sometimes a rapidly pulsed sound (a buzz) that had a
tonal quality was heard. The buzz was the only sound heard from
L. australis at any distance, and it was produced only occasionally;
consequently most attempts to listen to these porpoises were entirely
unsuccessful. No squeals (whistles) were heard; this was unex-
pected since we have heard squeals from other Lagenorhynchus
(L. albirostris, acutus, obliquidens, and the obscurus of New
Zealand).
These sounds of L. australis were low level and generally in-
audible beyond about 10-20 m. We estimate that the loudest
chcks were no more than -20 db re 1 dyne/cm- at 1 m, from
known hydrophone sensitivities and tape saturation levels, and
1971 PORPOISE SOUNDS 3
assumed supply voltages and amplifier gains. On only a few occa-
sions were we convinced that we knew which individual produced
the sounds that we heard, and therefore our estimates of signal
strength and of distance from the hydrophone are but guesses.
The click-sounds were of two types: a broadband click, and a
relatively restricted-bandwidth click (narrowband) at predomi-
nantly low frequencies. These two clicks never seemed to be
mixed. Both types were heard, we thought, from any one individ-
ual, with no obvious separation between the different kinds of
clicks, and no gradual transition. Though both types of clicks
were sometimes heard at slow rates (1 or 2 per sec), the broad-
band click was usually produced at a more rapid repetition-rate
(20 to 80 per sec.) than the narrowband click (5 to 25 per sec.).
The broadband click was shorter and had less energy at low fre-
quencies than the narrowband click. See the table for a comparison
of these two clicks.
The broadband click (Figs. lA and 2) was characterized by a
sharp onset, a short duration, as well as a more or less continuous
spectrum to 10 or 12 kHz, occasionally to 16 kHz. Analyses of
clicks showed a general drop in intensity of 1 to 2 db per 1000 Hz
above 5 or 6 kHz. This drop was greater than is consistent with
normal frequency-selective absorption for these distances and fre-
quencies, so we assume that this attenuation is characteristic of the
click of L. australis. The duration of the broadband click was
consistently a little less than 1 msec. Because of the general low
level of the sounds as well as their usual reduction in intensity at
higher frequencies, the clicks were easily masked by background
ambient.
The narrowband click (Figs. IB and 2) was restricted in fre-
quency to the lower 2000 Hz and appeared to have its greatest in-
tensity at or below 1000 Hz. Harmonics did exist, though at greatly
reduced levels. Analysis at high gain (but still undistorted) showed
some of the narrowband clicks with harmonic frequencies to 5 or
6 kHz. The narrowband click, with a duration of 1.5 to 3 msec,
usually occurred at slower repetition-rates (5 to 25 per sec), and
consistently had higher intensity at low frequencies than the broad-
band click. Perhaps because of its lower-frequency emphasis and
therefore better transmission characteristics, the narrowband click
was the one most commonly heard.
The third type of sound, the buzz (Fig. 3) was heard on a few
4 BREVIORA No. 366
occasions. This buzz had emphasis at discrete higher frequencies,
such that both the fundamental and high frequency overtones were
predominant in the aural impression of the sound. The buzzes
varied in duration from 0.6 to 1.1 sec. They were composed of a
pulsed fundamental near 300 Hz (Fig. 4) and strong overtones at
4 to 5 kHz. Two or three sidebands of the pulse repetition-rate
(modulation. Fig. IC) may be noted grouped around the 4- to
5-kHz overtone in spectral analysis (Fig. 3) of these buzzes (see
Watkins, 1967). The fundamental frequency of the buzz was more
intense than the overtones, yet at greater distances only the 4- to
5-kHz tone (with its associated sideband structure) was audible.
This was probably because of higher background ambient at the
lower frequencies. The buzz appeared to be produced at a higher
level than the clicks.
Discussion. Because of both the pulsed quality of the buzz and
its restricted frequency, we suppose that this sound was formed by
rapidly repeated narrowband clicks. Singly, the narrowband clicks
had few higher frequency components, but in a rapid series the
overtones were prominent. This is somewhat similar to sounds pro-
duced by Phocoena phocoena, composed of a rapid repetition of
narrowband clicks to form a continuous sound with selected higher
frequency emphases (Schevill, Watkins, and Ray, 1969). We did
not find the variation in the overtones of the buzz of L. australis
that we noted for Phocoena, but this may have been due to the
limited number of the former's buzz sounds that were recorded well
enough for such detailed analysis.
Perhaps the buzz was used in communication and it may have
been associated with stress. This could explain its relatively infre-
quent occurrence. The only time that the buzz was heard when
we thought we knew which porpoises were producing it (in Paso
Micalvi outside of Seno Grandi, Navarino Island, 6 December), a
group of three animals 15 to 20 m distant suddenly seemed to be
in a scuffle, darting at and away from each other. This sudden un-
usual activity coincided with the production of four buzzes, two of
them concurrently (Figs. 3 and 4), and so we assume that these
sounds were produced by these porpoises.
We have no evidence that Lagenorhynchus australis echolocates.
If the click sounds were used for echolocation as in some other
species ( Tursiops truncatus, Steno bredanensis, Orcinus orca,
Phocoena phocoena) , it must have been at relatively close ranges
1971 PORPOISE SOUNDS 5
because of the low level of the clicks. We did not hear any "ac-
celerando" in click series such as is typical of echolocation runs
during feeding by these other animals; however, we had no sugges-
tion that the L. aiistralis were feeding when the clicks were heard.
In fact, the clicks were not consistently associated with apparently
investigatory behavior by the animals. Porpoises sometimes passed
within a meter of the hydrophone and even appeared to return and
examine it without our detecting any sounds. On the other hand,
clicks were never heard unless a porpoise was close by.
The two click-types perhaps are equivalent to the two basic
click-categories noted for Tursiops by Norris, Evans, and Turner
(1967). They name these clicks by their function, ''discrimination
clicks" and "orientation clicks." The discrimination click of Tur-
siops has a reduced bandwidth and emphasis of lower frequencies,
while the orientation click has a wide bandwidth. In these respects
they match the sounds heard from L. australis, though no be-
havioral correlation was possible.
Although two types of clicks were heard, one with a relatively
restricted low frequency and other with broadband characteristics,
it suggests the possibility that only one click type exists in reality
and the variations noted result from changing orientation by an
animal possessing a directional sound system. Other cetaceans have
been shown to have such a directional sound field ( Tursiops, Nor-
ris, Prescott, Asa-Dorian, and Perkins, 1961; Orcinus, Schevill and
Watkins, 1966; Steno, Norris and Evans, 1967; and, Platanista,
Evans in Herald et al., 1969). Our data is insufficient to rule out
this possibility completely, but the evidence that we have seems to
argue instead for two distinct click types:
1 . The click durations of the two types are different. High fre-
quency emphasis in a low frequency click would not shorten the
length of the pulse but would simply extend the bandwidth.
2. The two click-types suddenly interchange with no pause be-
tween. We have no examples of a gradual shift from one type
to the other and we have very few individual clicks whose char-
acteristics are intermediate in form. Some of the subtle varia-
tions observed in the higher frequency components of successive
clicks of both types, however, may result from such direction-
ality, though we did not have opportunity to observe any cor-
relation of orientation with bandwith.
6 BREVIORA No. 366
Because of the difficulties we encountered in hearing the por-
poises, we were impressed with the low level of their sounds. We
also were acutely aware that it was not high background that
obscured their sounds, since the ambient noise levels in this region
were actually very low. Without carefully and recently calibrated
equipment, such low sound-levels are difficult (and probably mean-
ingless) to assess; however, our limitation much of the time ap-
peared to be the self-noise of the equipment rather than the local
ambient background. Perhaps the land barriers shielded the inland
channels from the usual open sea sounds and at the same time
provided enough shelter so that very little local wind and wave
noise was generated. In addition, we recognized very little contri-
bution of sound from other biological sources, and certainly these
porpoises had but small influence on the local ambient sound.
ACKNOWLEDGEMENTS
We thank the Antarctic Research Program of the National
Science Foundation for support and good help during the cruise
on board R/V HERO. The acoustic analysis and preparation of
this report were supported by the Office of Naval Research (Biol-
ogy branch) under contract Nonr 4446 and Nonr 241.09, and by
the National Science Foundation grant GA 1475. Experience and
observation were shared with the other members of the scientific
party aboard HERO; Kenneth S. Norris as chief scientist and
George Harvey were particularly involved and helpful. We thank
Elizabeth T. Bunce and Paul T. McElroy for criticism of the manu-
script.
LITERATURE CITED
Herald, E. S., R. L. Brownell, Jr., F. L. Frye, E. J. Morris, W. E. Evans,
AND A. B. Scott. 1969. Blind river dolphin: first side-swimming ceta-
cean. Science. 166 (3911): 1408-1410.
NoRRis, K. S., AND W. E. Evans. 1967. Directionality of echolocation clicks
in the rough-tooth porpoise, Steno hredanensis (Lesson). In W. N.
Tavolga (ed.), Marine Bio-Acoustics, vol. 2. Oxford, Pergamon Press.
Pp. 305-314.
Norris, K. S., W. E. Evans, and R. N. Turner. 1967. Echolocation in an
Atlantic bottlenose porpoise during discrimination. //; R.-G. Busnel
(ed.), Les Systemes Sonars Animaux, Biologic et Bionique, Jouy-en-
Josas, France. Pp. 409-437.
1971 PORPOISE SOUNDS 7
NoRRis, K. S.. J. H. Prescott, P. V. Asa-Dorian, and P. Perkins. 1961.
An experimental demonstration of echo-location behavior in the por-
poise, Tursiops truncatus (Montagu). Biological Bull.. 120 (2):
163-176.
SCHEVILL, W. E., AND W. A. Watkins. 1966. Sound structure and direc-
tionality in Orciniis (killer whale). Zoologica (N. Y.), 51 (2): 71-76.
ScHEViLL, W. E., W. A. Watkins, and C. Ray. 1969. Click structure in the
porpoise Phococna phocncna. Jour. Mammalogy. 50 (4): 721-728.
Watkins, W. A. 1963. Portable underwater recording system. Undersea
Technology, 4 (9): 23-24.
Watkins, W. A. 1967. The harmonic interval: fact or artifact in spectral
analysis of pulse trains. In W. N. Tavolga (ed.). Marine Bio- Acoustics,
vol. 2. Oxford. Pergamon Press. Pp. 15-42.
BREVIORA
No. 366
B
/ msec/div.
Figure 1. Oscillographic pictures of (A) the broadband click, (B) the
narrowband click, and (C) the pulse modulation of the buzz. Ambient
noise is superimposed on these sound traces.
1971
PORPOISE SOUNDS
8000-
Hz
4000-
I
0
'1
05
Seconds
Figure 2. Spectrographic analysis shows a burst of broadband clicks
followed by narrowband clicks. Although the latter become much greater
in amplitude as the animal conies closer, the frequency spectrum remains
relatively restricted. The bandwidth of the analyzing filter is 300 Hz. This
figure is the result of a repetitive analysis, with a small horizontal displace-
ment of the paper between analyses to widen artificially the traces of these
short-duration sounds for better photographic reproduction.
Narrowband click
Broadband click
Bandwidth
Principal frequency
Duration
Repetition rate
Intensity (re 1 dyne/cm2)
2 kHz
1 kHz or less
1.5 to 3 msec
5 to 25 /sec
— 20 db at principal
frequency
10 or 12 kHz
from less than 1 to 5 kHz
0.8 to 1 msec
20 to 80/sec
— 20db spread over
bandwidth
Table of characteristics of the two types of click
heard from Lagenorhynchiis australis.
10
BREVIORA
No. 366
7000-
5000-
Hz
2000-
Seconds
tw
i \
^ai
1.5
Figure 3. Two simultaneous buzzes have empiiasis in the 4- to 5-kHz
region as well as a strong fundamental at about 300 Hz. The analyzing
filter bandwidth is 300 Hz. Compare Fig. 4.
1000-
Hz
500-
■ ' ' Seconds
Figure 4. The fundamental frequencies of the same two buzzes shown
in Fig. 3 show variation in the region of 300 Hz. The continuous low fre-
quency band is ship's propulsion noise from the HERO about 5 or 6 miles
away. The analyzing filter bandwidth is 45 Hz.
BREVIORA
Mniseeim of Cojniiparative Zoology
Cambridge, Mass. 15 January, 1971 Number 367
MICROMISCHODUS SUGILLATUS, A NEW HEMIODONTID
CHARACIN FISH FROM BRAZIL, AND ITS RELATIONSHIP
TO THE CHILODONTIDAE
Tyson R. Roberts
Abstract. A new genus and species of Hemiodontidae, designated as
a new subfamily, Micromischodontinae, is described from the lower Rio
Negro and Middle Amazon of Brazil. Its osteology is described and
figured. It appears to be the hemiodontid genus most closely related to
Chilodontidae. The relationship between the functional and replacement
teeth of the pharyngeals suggests the probable manner in which the peculiar
pharyngeal teeth characteristic of Chilodontidae and Anostomidae evolved.
A brief definition is given of the family Chilodontidae.
ACKNOWLEDGEMENTS
Dr. Stanley H. Weitzman, Division of Fishes, U.S. National
Museum, suspected hemiodontids and chilodontids might be re-
lated and mentioned this to me two or three years ago. He is now
working on the relationships of these families and has graciously
let me examine unpublished illustrations of chilodontid osteology.
I am indebted to Sr. Heraldo Britski, curator of the fish collections
of the Museu de Zoologia, Universidade de Sao Paulo, and leader
of the Expediyao Permanente da Amazonia when the new fish was
collected, for permitting me to prepare its description. Professor
George S. Myers, Division of Systematic Biology, Stanford Uni-
versity. Dr. Weitzman and Sr. Britski reviewed the manuscript.
2 BREVIORA No. 367
INTRODUCTION
The hemiodontid herein described, collected by the Expedi^ao
Permanente da Amazonia' in 1967, represents a new subfamily.
While lacking certain morphological peculiarities of the highly
specialized Chilodontidae, it nevertheless appears to be more
closely related to them than is any other hemiodontid. In the
light of its discovery, there can be little doubt that Chilodontidae
and Hemiodontidae are indeed closely related. Although the
pharyngeal teeth of this new form are single cusped, the relation-
ship between functional and replacement teeth on the pharyngeals
suggests the primitive condition from which the peculiar multi-
cuspid pharyngeal teeth characteristic of Chilodontidae and
Anostomidae presumably evolved. Its highly distinctive trophic
structures indicate an unusual mode of feeding, perhaps similar to
that of Bivibranchia, considered to have the most specialized
trophic structures of all characoids. Nevertheless, it represents a
hne distant from Bivibranchia (and the related but less specialized
Argonectes) .
Hemiodontidae are marvelously streamlined, swift-swimming
fishes mostly six inches to a foot long. They form small groups in
open water in big rivers and lagos. Hemiodus are known in Brazil
as "voadores" because of their ability to jump. A group of
voadores leaping away from predaceous fishes or over a seine net
to safety is an impressive sight. Spawning presumably takes place
in open water and the young probably grow up in aggregations
staying near the bottom in fairly shallow open water. It appears
to be unrecorded whether hemiodontids form huge schools or
undertake major spawning migrations.
Hemiodontids hitherto known readily fall into two subfamilies,
Hemiodontinae and Bivibranchiinae, adults of which have multi-
cuspid teeth in the upper jaw and no teeth in the lower jaw. The
minute, unicuspid teeth and other features of the new fish differ so
greatly from previously known forms that it represents a new sub-
family.
1 The Expedicao Permanente da Amazonia, under the direction of Dr.
P. E. Vanzolini, is a cooperative effort among the Museu de Zoologia,
Universidade de Sao Paulo (MZUSP); Instituto Nacional de Pesquisas da
Amazonia (INPA) in Manaus; and Museu Goeldi in Belem. It is financed
by the Fundagao de Amparo a Pesquisa of the state of Sao Paulo. Much
attention has been devoted to fishes since fieldwork began in 1967. The
fish collections are housed at MZUSP.
1971 BRAZILIAN CHARACIN FISH 3
MICROMISCHODONTINAE, new subfamily
Highly streamlined, fusiform fishes typically hemiodontid in
habitus and osteology. Teeth pedicellate, with a single strongly
recurved cusp; tooth crowns black or brownish black, stalks deep
yellow or yellowish brown. Each dentary and lower pharyngeal
with two, nearly coextensive rows of 50-60 teeth. Teeth on upper
pharyngeal arranged in numerous, extremly regular rows (as in
Anostomidae). Upper jaw with a broad frenum, nonprotractile;
roof of mouth with fine, linear ridges, without valvelike structures.
Gill rakers elongate and numerous, with six to eight papillae on
either side of each gill raker, forming a dense carpetlike hning to
gill chambers. Lower pharyngeals exceptionally long and slender,
tooth-bearing for virtually their entire length. Upper limb of
second gill arch with a fleshy membrane forming a sort of pocket
anterior to upper pharyngeals. Posterior face of fourth gill arch
with ordinary gill filaments; face of fifth gill arch smooth; no dorsal
diverticulum between fourth and fifth arches. Stomach reversed,
that is, with cardiac portion (entrance of esophagus) posterior
and pyloric portion anterior in position; pyloric caeca about thirty,
not well differentiated; length of intestine in preserved specimens
about equal to standard length, forming a single loop upon leaving
stomach, then passing straight to vent. Posterior chamber of swim
bladder about one-half of standard length or six times length of
anterior chamber, terminating in a fine taper which extends to
above base of last anal fin ray. Adipose eye-fid thick and very
strong, extending from immediately behind nostrils well onto gill
cover, and with a narrow vertical sfit over the pupil (Fig. 1).
Cranial fontanels as in chilodontids and other hemiodontids,
anterior fontanel linear and narrow, posterior fontanel slightly
wider (Fig. 2). Size, shape, and position of jaw bones as in
Hemiodus; a peculiar fenestra in tooth-bearing portion of dentary
(a similarly located fenestra present in Hemiodus, absent in fore-
shortened dentary of chilodontids). Anterior end of ethmoid with
small lateral knobs. Circumorbital series with simple antorbital,
supraorbital, and full complement of six infraorbitals; first infra-
orbital smaller than those succeeding it (enlarged in Chilodon-
tidae), j^econd through fourth infraorbitals each slightly larger
than the preceding one. Branchiostegal rays five (five in Hemiodus
and Argonectes, four in Chilodus and Caenotropus); proximal end
4 BREVIORA No. 367
of fourth branchiostegal ray greatly expanded; hyoid bar general-
ized (apparently highly specialized in Chilodontidae). Gill mem-
branes free from isthmus, united to each other at a point below
middle of eye (broadly united to isthmus in chilodontids); isthmus
scaled anterior to cleithral symphysis (scaleless in chilodontids).
Three postcleithra; third (lowermost) postcleithrum with a lamel-
lar, posteriorly directed extension (as in Hemiodus). Weberian
apparatus and caudal skeleton without unusual modifications.
Forty vertebrae, including Weberian apparatus.
IVIICROMISCHODUS, new genus
Nomendatiiral type-species: M. sugillatus, new species
Body fusiform and highly streamlined. Secondary sexual di-
morphism unknown (specimens at hand collected in November
and December, with unripe gonads). Cranial roof smooth. Sides
of head largely covered by adipose eye-lids. Nares nontubular,
close-set and separated only by a flap flush with surface of head.
Tip of snout extends slightly beyond included lower jaw. With
mouth fully opened, gape almost vertical and about as large as
eye diameter. With mouth closed, dorsoposterior edge of maxillary
slips under first infraorbital bone; maxillary not extending as far
back as anterior margin of eye, but only to below posterior nostril.
Toothless portion of lower jaw (posterior to rictal membrane)
about four or five times longer than tooth-bearing portion. Articu-
lation of lower jaw below middle of eye; rictal membrane below
nostril, distinctly in front of anterior orbital rim. Lateral line
complete, slightly decurved anteriorly, then running just below
lateral midline of body to last scale row on caudal base.
Origin of dorsal fin midway between snout tip and base of
caudal fin. Anal fin small. Caudal fin deeply forked. Dorsal, anal,
and median caudal fin rays Vv'ith well-developed, overlapping, mem-
branous lappets or alae (Fig. 1 ). Similar structures occur in many
fast-swimming lower teleosts, including various other characoids,
some cyprinoids and clupeoids, and Chanos.
Known by a single species from the Middle Amazon and lower
Rio Negro in Brazil.
Derivation of name. Micromischodus is from the Greek micro,
small, little; mischos, stalk, petiole, peduncle; and odous (odon),
odontos, masc, tooth.
1971 BRAZILIAN CHARACIN FISH 5
M. SUGILLATUS, new species
Figure 1
Note. Standard lengths are used throughout this paper. Un-
less stated otherwise, proportional measurements are expressed
as times in standard length. Counts and measurements are given
for the holotype first, followed in parentheses by the ranges for
the paratypes.
Holotype. MZUSP 6773, 152.0 mm, Igarape Tarumazinho,
15 km NW of Manaus, on the left bank of the Rio Negro,
Estado do Amazonas, 17-18 November 1967.
Paratypes. MZUSP 8870, six specimens 125.0-144.6 mm,
same data as holotype; MZUSP 6691, four specimens 107.3-137.1
mm, Rio Negro, arredores de Manaus, Estado do Amazonas,
15-23 November 1967; MCZ 46718, two specimens 118.4 and
137.1 mm, Rio Canuma, Canuma (near Maues), Estado do
Amazonas, 28-29 November 1967; MCZ 46719, seven specimens
79.5-108.9 mm, Igarape do Rio Jamari, Terra Santa, Estado do
Para, 14 December 1967.
Proportional measurements. Greatest body depth (at origin of
dorsal fin) 4.1 (4.0-4.5). Greatest body width (below dorsal
fin) 5.7 (5.6-6.6). Least depth of caudal peduncle 11.4 (11.3-
12.4). Distance from snout-tip to dorsal origin 2.01 (1.94-2.08).
Length of caudal peduncle 7.45 (6.4-7.8). Head 3.4 (3.1-3.4).
Eye 3.9 (3.4—4.1) in head. Bony interorbital space 3.8 (3.5-4.1)
in head. Snout 3.5 (3.4-3.8) in head. Width of mouth (measured
to outer sides of maxillaries) 4.1 (4.1-4.7).
Fins. Fins, excepting caudal, scaleless. Dorsal fin rays 11
(11 in all paratypes except one with 10), first two rays simple,
last ray divided to its base. Proximal two-thirds to three-quarters
of each ray with well-developed membranous lappets or alae,
largely overlapping when fin is depressed. Dorsal fin margin
falcate. Height of dorsal fin 4.7 (4.3-4.8). Base of dorsal fin 9.2
(8.1-10.3). Anal fin small, about one-third as large as dorsal fin,
with similar membranous lappets. Anal rays 10 (10), first two
rays simple, last ray divided to its base. Height of anal fin 8.2
(7.4-8.3). Anal fin margin falcate. Base of anal fin 12.4 (11.8-
13.4). Caudal fin deeply forked. Lower caudal lobe slightly
larger than upper caudal lobe, with its upper margin shghtly con-
vex (Fig. 1). The lower caudal lobe is similarly modified in other
BREVIORA
No. 367
00
B
4)
c
a
B
B
q
CO
3
u
a
■4-*
45
en
a
60
s
"n
en
a
o
tn
s
o
u
3
oo
1971 BRAZILIAN CHARACIN FISH 7
fast-swimming characoids, for example in Hydrocynus goliath
(Boulenger, 1899, pi. 11). Principal caudal rays 10 -f- 9. Median
four to six rays with well-developed lappets. Procurrent rays
moderately numerous (9 + 7 in 125-mm specimen), unspecial-
ized. Adipose fin small, its base over posterior portion of anal
fin base. Pectoral and pelvic fins with two or three peculiarly
thickened and elongated scales forming a sheath for edge of outer-
most rays. Pectoral fin rays 21 (18-22). Length of pectoral fin
6.3 (5.3-6.5). Pelvic fin rays 11 (11), outermost ray only un-
branched. Length of pelvic fin 7.0 (6.1-7.2).
Scales. Scales cycloid, or with laciniate (but not ctenoid)
borders. Body completely scaled, 57 (56-64) scales in a lateral
series; about 21 (16-22) predorsal scales (irregularly aligned);
11 (11 or 12) scales between lateral line and dorsal origin; 4 (4)
scales between lateral line and pelvic insertion; 20 (17-22) scales
between last dorsal ray and adipose fin; 15 (14-16) scales along
midline between symphysis of cleithra and pelvic base; and 16
(16-18) scales around caudal peduncle. Prepelvic scales below
lateral line gradually increasing in size ventralwards; scales on
abdomen about two or three times larger in diameter than scales
above lateral line. Caudal fin beyond hypural fan with two or
three rows of scales.
Coloration in alcohol. After two years in preservative (fixed in
formalin while alive and transferred to ethyl alcohol within a few
months) the specimens are straw colored, darkest above. The
most notable feature is the solid black or blue-black color on the
membranes between the distal halves of all of the dorsal fin rays
(but not on the rays themselves), (The hemiodontid Argonectes
scapularis and chilodontid Tylobronchiis maculosus have similarly
marked dorsal fins. ) Pectoral, pelvic, anal, and adipose fins color-
less; caudal fin dusky, lower lobe of caudal becoming darker (but
not as dark as dorsal fin) near its tip. A faint longitudinal stripe
running length of the body but stopping at shoulder, not running
through eyes or onto caudal fin. Above lateral line fine melano-
phores lie beneath the scales and in concentrations paralleling
exposed scale margins. Below lateral line melanophores almost
entirely absent in some specimens; in other specimens melano-
phores are virtually absent beneath the scales but are present
along the scale margins. The reticulated pattern of vertically
oriented spots thus produced is similar to the pattern present in
8 BREVIORA No. 367
all Chilodontidae, except that since the scales are much smaller,
the spots are small and not as dark. In all specimens body dis-
tinctly lighter below lateral line than above. Gill cover marked
by an indistinct dark oval patch caused by a concentration of
melanophores in the membrane lining inside of gill cover, which
shows through the opercular bones.
The trivial name sugillatus (Latin: sugillo, -atus, beaten black
and blue; sugiUatum, black and blue spot, bruise) refers to the
coloration on the dorsal fin and opercles.
Ecological notes. According to Sr. Britski, specimens of M.
sugillatus were collected by seining along shore in lakes or places
with slow-flowing, black or crystalline water. A very large number
of species, predominantly characins, was obtained at these locali-
ties. The stomach contents of specimens from the Igarape do
Rio Jamari include an assortment of bottom material, including
many of what appear to be droppings of small fishes. Very small
insects are well represented, including larval Diptera and, most
abundant, a corixid (identified by Dr. John Lawrence) about L5
mm long. These are about the largest organisms in the stomach
contents.
OSTEOLOGY
Figures 2—15
Illustrations of the osteology of M. sugillatus are based on a
125.0-mm paratype from MZUSP 8870. Supplemental observa-
tions were made on a 102.2-mm paratype from MCZ 46719. The
nomenclature of bones follows Weitzman (1962) except that
"vomer" and "intercalar" are used in place of "prevomer" and
"opisthotic."
Cranium (Figs. 2-4). Roofing bones of skull smooth and flat-
tened. Cranial fontanels narrow, complete. Anterior fontanel
linear and about half as wide as posterior fontanel for almost its
entire length, but widening immediately in front of epiphyseal bar.
Ethmoid narrow, with two lateral knobs anteriorly articulating with
either premaxillary, ventro-lateral laminar projections immediately
posterior to these knobs, and a median cleft in its posterior half
(Fig. 2). Supraoccipital crest flat, not extending as far as end of
cranium. Posttemporal fossae well developed. Dilator groove well
developed, frontal participating in its formation; dorsal Umit of
1971
BRAZILIAN CHARACIN FISH
antorbjtal
nasa
vomer
ethmoid
parietal
epiphyseal bar
lateral ethmoid
frontal
sphenotic
epiotic
exoccipital
supraoccipital
premaxillary
infraorbital 6
Dterotic
Figure 2. Cranium of Micromischodiis siigillatiis (dorsal view),
osteological figures based on 125-mm specimen from MZUSP 8870.
All
dilator groove marked by a concavity in dorsolateral margin of
frontal bone; sphenotic spine lamellar, rounded at tip.
Ethmoid separated from vomer by a cartilaginous septum (in-
dicated by heavy stippling in Figure 3). Vomer with peculiar
anterior knoblike processes on either side. Lateral ethmoid rela-
tively large, with elongate, strutlike process articulating with knob-
hke process of vomer (Figs. 3 and 4) and a small median projec-
tion firmly articulated to antero-ventral comer of rhinosphenoid.
Rhinosphenoid large, sharing strongly interdigitating sutures with
antero-ventral projection of orbitosphenoid (Fig. 3). Cartilaginous
septum between dorsal edge of rhinosphenoid and roof of cranium
and another between ventral margin of rhinosphenoid and para-
sphenoid (cartilage indicated by heavy stippling in Figure 3).
Rhinosphenoid and orbitosphenoid not directly contacting para-
sphenoid. Parasphenoid deeply cleft posteriorly, the thin divisions
thus formed cleft at their tips (Fig. 4; compare with Weitzman,
1962, fig. 4 of Brycon on p. 60; and Roberts, 1969, fig. 20 of
Acestrorhynchus on p. 463). Intercalar well developed. Pterotic
with strong posteriorly directed spine.
Jaws and jaw suspension (Figs. 2, 5-7). The size and shape
of the jaw bones and their relationships to each other are similar
10
BREVIORA
No. 367
lateral ethmoid ^"^""^^l ^Phenotic pterotjc parietal supraocclpital
ethmoid
vomer
rhinosphenoid
orbitosphenoid
pterosphenold parasphenoid Prootic '"<ercalar "basioccipital
Figure 3. Cranium of Microinischodus sugillatiis (lateral view).
to those in Hemiodus. PremaxiUary with a slender, medially
directed symphyseal process, round in cross section and blunt at
the tip, which approximates but does not articulate or contact its
opposite fellow. Lateral surface of premaxillary with a shelflike
projection posterior to which is a groovelike depression into which
snugly fits anterodorsal corner of maxillary. Dorsal portion of
parasphenoid
pterosphenold
mesopterygoid
palatine gctopterygoid
hyomandibular
epiotic
basioccipital
ethmoid
vomer
exoccipital
lateral ethmoid
frontal
orbitosphenoid
intercalar
pterotic
sp hen otic
prootic
Figure 4. Cranium of Miciomischodus sugillatiis (ventral view).
1971
BRAZILIAN CHARACIN FISH
11
premaxillary and maxillary firmly embedded in a tough connective
tissue that binds them to tip of ethmoid although they do not con-
tact it. Maxillary with dorsoposterior corner expanded and con-
nected by a broad, strong ligament to palatine (Fig. 2). Maxillary
bearing a short segment of cephalic sensory canal (Fig. 5).
Dentary with a slender, median symphyseal process, round in
cross section, blunt at the tip which approximates and is strongly
joined by ligaments to its opposite fellow. Tooth-bearing portion
of dentary elevated; rictal membrane attaches maxillary to lateral
surface of elevated portion of dentary. Portion of dentary pos-
terior to gape gradually diminishing in width, about twice as long
as portion anterior to rictus; articulation with quadrate beneath
infraorbitals
supraorbital
antorbital
premaxillary
ope re I e
subopercle
maxillary
interopercle
preopercle
quadrate
angular
articular
dentary
Figure 5. Jaws and facial bones of Micromischodiis siigiUatiis (lateral
view).
middle of eye. Elevated portion of dentary with a peculiar fenestra
below part of the tooth rows (Figs. 5-7); a fenestra identical in
shape and position occurs in the toothless dentary of Hemiodus
but is lacking in chilodontids.
Premaxillary with about 14 minute moveable teeth. Maxillary
toothless. Dentary with two coextensive rows of minute moveable
teeth. The 125-mm specimen has about 54 teeth in outer row
and about 60 (slightly smaller) teeth in inner row, uppermost ten
12
BREVIORA
No. 367
of which are nonpedicellate; replacement teeth much in evidence
(Fig. 7).
The palatine arch presents no unusual features. Metapterygoid-
quadrate foramen well developed. Hyomandibular well separated
from mesopterygoid.
Facial bones (Figs. 5-6). Facial bones present no unusual
features. Circumorbital series complete. Nasal bone tubular,
laminar projections minute. Antorbital slender, struthke. Infra-
orbitals six; infraorbitals 2-4 gradually increasing in size over ones
preceding them. Infraortibal 1 not enlarged. Infraorbitals 3 and
4 with tubes from postorbital branch of cephalic sensory canal
system leading to their posterior margin (Fig. 5).
Posterior bony margin of gill cover formed largely by sub-
opercle; subopercle with well-developed, dorsally-projecting spine
at anteroventral end. Preopercle with a slender tubular extension
dorsally. Suprapreopercle absent.
Visceral arches (Figs. 8-10). Hyoid arch without unusual fea-
tures. Branchiostegal rays five (as in Hemiodus, Argonectes),
first four articulating with ceratohyal, fifth with epihyal. Proxunal
end of fourth where it connects with ceratohyal greatly expanded
(see Fig. 9).
premaxillary
maxillary
palatine
dentary
coronomeckelian bone
opercle
subopercle
articular
Interopercle
preopercle
Figure 6. Jaws, jaw suspension, and opercular bones of Micromischodus
siigillatus (internal view).
1971
BRAZILIAN CHARACIN FISH
13
Figure 7. Tooth-bearing portion of dentary of Micromiscliodiis
sugillattis (internal view).
14
BREVIORA
No. 367
ventral liypohyal
dorsal hypohyal
branch lostegal rays
Figure 8. Hyoid arch, branchiostegal rays, and urohyal of Micro-
mischodits siigillatiis (dorsal view).
Pharyngeal teeth (Figs. 9 and 10). Lower phaiyngeals pecu-
liarly elongate and bearing two rows of teeth for virtually their
entire length. The recurved crowns of the teeth in anterior row on
lower pharyngeal project forward; replacement teeth lie anteriorly
to their bases (Fig. lOA). On the trailing edge of the lower
pharyngeals is a row of smaller teeth with the recurved crowns
projecting backwards; replacement teeth lie posterior to their
bases (Fig. lOB). In 125-mm specimen about 64 teeth in anterior
row and 60 in posterior row.
At first glance the anterior and posterior rows of lower pharyn-
geal teeth each appear to constitute two rows. This is because
1971
BRAZILIAN CHARACIN FISH
15
basihyal
basibranchials
hypobranchials
ceratobranchials
epibranchials
suspensory pharyngeals
upper pharyngeals
lower pharyngeal
Figure 9. Gill arches of Micromischodus sitgillatiis (dorsal view). Left
side in normal position, right side with uppermost elements pinned out and
to the side.
not all of the teeth have assumed the "definitive" functional posi-
tion; teeth just coming into position are out of line with the rest.
Since none of the teeth are ankylosed to the bone, the borderline
between teeth in functional position and replacement teeth is
indistinct. Evidently some of the replacement teeth are "func-
tional" before they are in the definitive functional position, and
even before the functional teeth they are about to replace have
fallen out. Tooth replacement is much in evidence, and probably
occurs in waves proceeding the length of the lower pharyngeals.
In some places a functional tooth may have two replacement teeth
at its base (see Fig. lOB). These observed characteristics are
16
BREVIORA
No. 367
highly suggestive of the sort of situation from which the pedicellate
multicuspid pharyngeal teeth of Chilodontidae and Anostomidae
presumably evolved. If this view is correct, then successive cusps
fused below the main cusp to the stalklike base are phylogenetic-
ally equivalent to successive generations of replacement teeth.
Figure 10. Lower pharyngeal teeth of Micromischodiis sugillatiis (pos-
terior view). A, anterior row; B, posterior row.
The upper pharyngeals bear numerous regularly arranged rows
of teeth in which the crowns project backwards (in opposition to
crowns of teeth in anterior row of lower pharyngeal which point
forwards). The third suspensory pharyngeal bears a small patch
of similar teeth. Gill rakers edentulous.
The pharyngeal teeth, although morphologically similar to those
in the jaws, are more solidly implanted and much larger (except
for the teeth in the posterior row on lower pharyngeal); still, they
are considerably smaller than the greatly enlarged and very solidly
implanted pharyngeal teeth of Chilodontidae and Anostomidae.
In the 125-mm specimen the teeth in the upper jaw are about 0.2
mm long; the teeth in the outer row of the lower jaw are 0.4 mm
long; the teeth on the upper pharyngeal and in the anterior row on
1971
BRAZILIAN CHARACIN FISH
17
the lower phaiyngeal are 1.0 mm long; and the teeth in the pos-
terior row on the lower pharyngeal are 0.4 mm long. The highly
regular arrangement of tooth rows on the upper pharyngeal is
suggestive of the regular rows on the upper pharyngeals of An-
ostomidae. The arrangement of teeth in the lower pharyngeals is
unique. In most characoids, including Hemiodus, the teeth are
in several irregular rows on an expanded portion of the lower
pharyngeal near the midline; the distal half of the lower pharyngeal
is usually toothless.
Weberian apparatus (Figs. 11 and 12). Weberian apparatus
showing no unusual features, similar to that of typical Characidae.
Neural arch pedicle of third vertebra with a dorsally directed pro-
cess tightly bound into a groovehke fossa in base of neural arch
complex. Base of neural arch complex extends posteriorly so that
its posteroventral surface almost contacts entire anterior face of
neural spine of fourth vertebra. Rib of fifth vertebra with a
medially directed process ligamentously attached to os suspen-
sorium. Transverse process of third vertebra, which cradles inter-
calarium, well developed. Lateral process of os suspensorium very
neural complex
claustrum
neural arch of third vertebra
transverse process of third vertebra
supraneural
neural arch and spine
of fourth vertebra
scaphium
intercalarium
pleural rib
lateral process of second/
vertebra tripus'
OS suspensorium
Figure 11. Weberian apparatus of Micromischodus sugillatus (lateral
view).
18
BREVIORA
No. 367
neural complex
scaphium
lateral process of centrum 2
tripus
rib of fifth vertebra
OS suspensorium
centrum 4
centrum 5
Figure 12. Weberian apparatus of Micromischodus sugillatiis (ventral
view).
large, overlying transformator process of tripus. Neural arch of
third vertebra extending far anteriorly over second centrum (Fig.
11). Lateral process of centrum 2 very large.
Pectoral girdle (Fig. 13). Pectoral girdle without unusual
specializations. Three postcleithra. Most distinctive feature is
lamellar projection from proximal part of third postcleithra (simi-
lar process present in Hemiodus); in other characoids third post-
cleithrum slender for entire length.
Pelvic girdle (Fig. 14). Ischiac process well developed; four
radials; pelvic splint present; rays invariably 11.
Caudal skeleton (Fig. 15). Caudal skeleton generahzed. Hy-
purals 5 and 6 fused, rest separate. Two uroneurals. Three
epurals. Principal caudal rays invariably 10 + 9. Nine upper
and seven lower procurrent rays in 125-mm specimen.
'
197
BRAZILIAN CHARACIN FISH
19
posttemporal
supracleithrum
postcleithra
extrascapular
coracoid
Figure 13. Pectoral girdle of Micromischodus sugillatus (internal view).
Inset: external view of extrascapular and posttemporal.
20
BREVIORA
No. 367
radials
ischiac process
Figure 14. Pelvic girdle of Micromischodus sugillatus (ventral view).
Enlarged view on right side with radials exposed by removal of outer half
of each lepidotrich.
1971
BRAZILIAN CHARACIN FISH
21
upper procurrent rays
epurals
uroneural 2
hypural I
hypural 2
uroneural I
neural spines
haemal spines
upper principal rays
hypural centrum
hypural 7
lower principal rays
lower procurrent rays'
Figure 15. Caudal skeleton of Micromischodus sugillatiis (lateral view).
22 BREVIORA No. 367
DEFINITION OF THE FAMILY CHILODONTIDAE
A definition of the family Chilodontidae is given here primarly
for comparison with Micromischodontinae. This definition is not
the result of thorough researches of all taxa involved; it is an-
ticipated that studies in progress by Weitzman will gready extend
knowledge of the group. Trophic structures were emphasized in
defining Micromischodontinae, as they will also be in the following
definition. Modification of trophic structures has been the main
theme in the great adaptive radiation of characoids. Trenchant
differences in feeding habits and trophic structures characterize
most genera and higher taxa, and it is natural to emphasize these
differences in definitions. Some important modifications which do
not directly involve trophic structures are nevertheless strongly
linked to modes of feeding, viz., swimming position in Chilo-
dontidae and Anostomidae. Parenthetically, any attempt to work
out a phyletic classification of the higher characoid taxa must
necessarily give major consideration to the evolution of their
trophic structures.
Chilodontidae. Characoid fishes reaching about 150 mm in
standard length which spend much of the time in an oblique head-
down swimming position. In many regards — including position
of fins, approximate number of fin rays, and general cranial osteol-
ogy— they are like Hemiodontidae. They differ from the Hemi-
odontidae in having the skull, including the lower jaw, and the
whole body relatively short, and in their trophic modifications.
Number of scales considerably less than in Hemiodontidae (about
25-30 vs. 50-125). Pectoral fin with 13-16 rays (18-23 in
Hemiodontidae). Vertebrae including Weberian apparatus 33 in
Chilodus, 35 in Tylobronchus (Eigenmann, 1912: 271-273).
Lower jaw behind rictal membrane greatly foreshortened; first
infraorbital enlarged; jaw teeth and pharyngeal teeth reduced in
number. Premaxillary with a single row of conical {Caenotropus,
Chilodus) or bicuspid {Tylobronchus), stalked teeth; lower jaw
with a single row of conical, stalked teeth {Chilodus, Tylo-
bronchus) or no teeth in large adults {Caenotropus). Teeth em-
bedded in lips and freely moveable, not ankylosed to jaw bone.
Upper and lower pharyngeals with enlarged teeth with two, three,
or more cusps ossified to a stout, stalklike base. In Chilodus,
lower pharyngeal teeth confined to a small, round patch in the
1971 BRAZILIAN CHARACIN FISH 23
center of a large, bowllike depression formed by concave dorsal
surfaces of the greatly expanded (and highly peculiar) lower
pharyngeal bones. Posterior face of fourth arch and exposed face
of fifth arch bearing complementary rosettes of what appear to be
modified gill filaments covered with tiny papillae and with tough
booklets at their distal ends, evidently adapted to food selection
(see Gery, 1964, fig. 2 on p. 63). Gill rakers reduced in number;
bony lamellae of gill rakers reduced or absent in Chilodiis (in-
variably well developed in Hemiodontidae). Gill membranes
tightly bound to posteriormost part of isthmus, i.e., immediately
in front of base of pectoral girdle (free in Hemiodontidae). Hyoid
bar highly modified. Branchiostegal rays four (verified in Caeno-
tropiis and Chilodus). Isthmus scaleless. Caudal fin less forked
and anal fin larger than in Hemiodontidae. Membranous lappets
on dorsal, anal, and caudal fins reduced. Adipose eyelid reduced.
Distal half of dorsal fin wholly or partly covered with a black or
bluish black color, which partly breaks up into spots in Chilodus;
sides of body with a punctuate color pattern, spots borne on in-
dividual scales, and usually with a dark, longitudinal stripe on
the midline extending through the eye onto the middle of the caudal
fin.
1 have pointed out (Roberts, 1969: 424, 442) that Chilo-
dontidae and Anostomidae have similar multicuspid pharyngeal
teeth, and suggested that perhaps the Chilodontidae should be
regarded as a subfamily of Anostomidae. In the light of the present
study, I still feel that the two groups are closely related, but am
inclined to regard Chilodontidae as of family rank. The chilo-
dontids are set oflf from Anostomidae by the specialized structure
of their fourth and fifth gill arches and form a small group of
clearly related forms. And while it seems clear that the chilo-
dontids are related to the Anostomidae, the anostomid genus
closest to them cannot be singled out. Furthermore, the chilo-
dontids show equally clearly indications of relationship with
Hemiodontidae — a relationship closer than I previously had
thought likely. Thus the reasonable solution for now, and one
which may prove of lasting value, is recognition of three families,
Hemiodontidae, Chilodontidae, and Anostomidae. It will be noted
that the Anostomidae form a "well-knit" group of some ten or
eleven genera.
24 BREVIORA No. 367
COMMENTS ON HEMIODONTINAE
AND BIVIBRANCHIINAE
The Hemiodontinae comprises Hemiodus (but see Gery, 1963),
with numerous species differing sharply in color patterns and scale
counts (see Bohkle, 1955) and two or three closely related genera
of doubtful distinctness and confusing nomenclatural status (see
Gery, 1961; 1963). The Bivibranchiinae comprises three quite
distinct genera, each with but one or two species, Argonectes
(Bohlke and Myers, 1956), Atomaster (Eigenmann and Myers,
1927) and Bivibranchia. Briefly, the Bivibranchiinae have a highly
protractile upper jaw with tricuspid teeth, and highly specialized
trophic structures in the roof of the mouth and in the pharynx.
The nature of their pharyngeal teeth has yet to be elucidated. Of
the three genera, Argonectes is least specialized and approaches in
some respects the Hemiodontinae, which have relatively nonpro-
tractile upper jaws with multicuspid teeth of eight or nine cusps,
and relatively generalized oral and pharyngeal passages.
In neither of the two subfamiUes do adults have teeth in the
lower jaw. The dentition of young Hemiodus, however, is very
different from that of adults. According to Gery (1963: 604), in
alevins of Hemiodus (tentatively identified as H. unimaculatus)
up to 17.3 mm in standard length, there are but six to eight teeth
on the premaxillaries, these teeth having fewer cusps than those
in adults, and the dentary bears four or six very small conical teeth,
visible only with strong magnification. We may feel fairly secure
in supposing (Roberts, 1967) that the earliest dentition in
Hemiodus consists of small conical teeth in both upper and lower
jaws. As Hemiodus grow, presumably the upper jaw teeth increase
in number and are replaced by teeth with successively more cusps,
whereas the conical teeth in the lower jaw are replaced relatively
few times, if at all, and drop out altogether at a fairly early stage.
Menezes and Oliveira e Silva (1949) reported that stomach
contents of Hemiodus parnaguae from the Rio Parnaiba in Piaui,
Brazil, contain mud, algae, and remains of higher plants. No in-
formation is available on food habits of Bivibranchiinae. I suspect
that Bivibranchia take a mouthful of sand, sort food particles out
in the gill chambers and then spit the sand out of the mouth.
Gery (1969: 836) stated that Bivibranchia bury themselves in
sand "like sand-eels" but gave no further details. Possibly this
observation involves a feeding activity.
1971 BRAZILIAN CHARACIN FISH 25
LITERATURE CITED
BoHLKE, J. E. 1955. Studies on fishes of the family Characidae. — No.
lU. Notes on the coloration of the species of Hemiodiis, Pterohemiodiis
and Aiiisitsia, with the description of a new Hemiodiis from the Rio
Negro at the Brazil-Colombia border. Acad. Nat. Sci. Philadelphia,
Notulae Naturae, No. 278: 1-15.
, AND G. S. Myers. 1956. Studies on fishes of the family
Characidae. — No. 11. A new genus and species of hemiodontins from
the Rio Orinoco in Venezuela. Acad. Nat. Sci. Philadelphia, Notulae
Naturae, No. 286: 1-6.
BouLENGER, G. A. 1899. Materiaux pour la Faune du Congo. Poissons
nouveaux du Congo. Ann. Mus. Congo (Tervuren), Zool., ser. 1,
1 (pts. 1-6): 1-164.
EiGENMANN, C. H. 1912. The fresh-water fishes of British Guiana.
Mem. Carnegie Mus., 5: xxii -\- 578 pp.
, AND G. S. Myers. 1927. A new genus of Brazilian characin
fishes allied to Bivibranchia. Proc. Nat'l. Acad. Sci. (Washington),
3(8): 565-566.
Gery, J. 1961. Pterohemiodiis liielingi sp. nov.. un curieux poisson
characoide a nageoire dorsale filamenteuse, avec une cle des genres
d'Hemiodontinae (Ostariophysi-Erythrinidae). Bonner Zoologische
Beitrage. 314(12): 332-342.
. 1963. Sur la nomenclature et la systematique du genre
Hemiodiis Miiller (Pisces, Characoidei). Bull. Mus. Nat'l. Hist. Nat.,
ser. 2, 35(6): 598-605.
1964. A review of the Chilodinae, with a key to the
species. Tropical Fish Hobbyist, May, 1964: 5-10, 63-67.
1969. The fresh-water fishes of South America. In Fitlkau,
E. J., et alia (editors), Biogeography and Ecology in South America.
Vol. 2, pp. 828-848. The Hague, Junk.
Menezes, R. S. and S. L. Oliveira e Silva. 1949. Alimentagao de
voador, "Hemiodus parnaguae" Eigenmann and Henn, da bacia do Rio
Parnaiba, Piaui. Rev. Bras. Biol., 9(2): 241-245.
Roberts, T. R. 1967. Tooth formation and replacement in characoid
fishes. Stanford Ichth. Bull., 8(4): 251-259.
1969. Osteology and relationships of characoid fishes, par-
ticularly the genera Hepsetiis, Salminiis, Hoplias, Ctenoliiciiis, and
Acestrorhynchus. Proc. California Acad. Sci., ser 4, 36(15): 391-500.
Weitzman, S. H. 1962. The osteology of Brycon meeki, a generalized
characid fish, with an osteological definition of the subfamily. Stan-
ford Ichth. Bull., 8(1): 1-77.
BREVIORA
Museium of Compsirative Zoology
Cambridge, Mass. 29 January, 1971 Number 368
STRUCTURAL HABITATS OF WEST INDIAN ANOLIS
LIZARDS I. LOWLAND JAMAICA
Thomas W. and Amy Schoener^
Abstract. This paper reports differences in structural and climatic
habitat among the commonest Anolis species of three lowland Jamaican
localities. Three of the species studied, grahami, opalinus, and valencienni,
occurred at all localities; two other species, sagrei and Uneatopiis, occurred
at one (Whitehouse) and two (Port Antonio, Mona) localities, respec-
tively, and thus were complementary.
With a few exceptions, the ordering of species by height, from highest
to lowest, was valencienni, grahami, opalinus, and Uneatopiis or sagrei; the
ordering by diameter, from thickest to thinnest, was opalinus, grahami,
lineatopus or sagrei, and valencienni; the ordering by insolation, from
sunniest to shadiest, was sagrei, grahami, valencienni, lineatopus, and
opalinus; the ordering by size, from largest to smallest, was valencienni,
lineatopus, grahami, sagrei, and opalinus. Within species, larger individuals
tended to occur higher and on thicker perches, smaller individuals lower
and on thinner perches.
The above orderings result in low interspecific spatial overlap of similarly
sized individuals in two ways. First, a direct relationship between body size
and perch diameter within species and an inverse one between species
ensures that spatially abutting species will overlap most their respective
individuals least alike in size. Second, a direct relationship within species
between size and height also, except for valencienni and grahami, results
in the greatest spatial overlap being between the most dissimilarly sized
individuals.
Many statistically significant associations were found between the habitat
and climatic variables; the most common was a tendency for thin perches
to be more often occupied in the sun.
1 Biological Laboratories and Museum of Comparative Zoology, Harvard
University, Cambridge, Mass. 02138.
2 BREVIORA No. 368
This is the first of a series of papers describing in a standard
way the structural habitat of some West Indian Anolis Hzards. Its
primary purpose is to document differences between the size and
sex classes of all the Anolis species found within a particular,
limited study area. Between-site comparisons will be drawn occa-
sionally; however, because of the great intersite variation in vege-
tation structure and its effect on lizard habitat distributions, that
aspect will be described and analyzed in detail in a larger work
covering much of the western Caribbean.
The "structural habitat," a term first invented by Rand (1964)
for Anolis, refers to the spatial niche of a species population de-
scribed in terms of characteristics of the vegetation and other
structures upon which these arboreal animals carry out their activi-
ties. The two such characteristics that Rand originally used and
that have subsequently proven most useful in discriminating the
various species or species-classes are perch height and perch
diameter (Rand, I.e., 1967a; Rand and Rand, 1966; Schoener,
1968; Schoener and Gorman, 1968; Laska, 1970), and these will
be the ones used below. Other possible structural characteristics
are perch texture, perch color, and the size and nature of the plant
or other object to which the perch is attached.
Observations that combine to give an overall picture of the
spatial dimensions of the niche must be summed over a given
time period and over a particular set of individuals. The method
chosen in this and succeeding studies is to lump together "first
sightings" recorded continuously throughout the day or throughout
the period of maximum activity from a population of several hun-
dred animals for each of the the commonest species. Thus, ideally,
the sites of the major activities — thermoregulation, searching for
food, capturing and eating prey, and social interaction — are
weighted when observations are combined according to that frac-
tion of the day spent by the average individual in the particular
activity.
There are several hmitations and qualifications of the structural-
habitat concept which must be mentioned at the outset.
First, it does not provide a true picture of the animals' daily
perch distribution, because inconspicuous animals are more often
missed. This means that the most visible activities, feeding or fight-
ing, for example, are disproportionately weighted in the totals. In-
conspicuousness is perhaps less of a problem for continuous
observation of single individuals than for "first sightings."
1971 JAMAICAN ANGLES 3
Second, weighting activities in proportion to the amount of time
they take will not necessarily produce the most meaningful measure
of a species' spatial distribution, either for testing hypotheses of
optimal individual behavior or of community composition. For
example, it may be most useful to know the places where prey are
captured in order to determine how similar species can be and still
coexist, but this activity consumes a very small amount of time, and
Andrews (1971) and Trivers (personal communication) have
both shown that distributions of feeding sites can differ markedly
from the overall structural habitat. However, taxonomic break-
down of the prey of the four Bimini Anolis has shown that the
prey is qualitatively what would be predicted from the overall
structural habitat (Schoener, 1968).
Third, consideration of structural habitat alone provides an in-
complete picture of the total niche of a species. The most impor-
tant other kind of property characterizing the space in which an
anole lives is climatic; some measure of this is given below for
certain situations. A second important way in which animals of
the same structural habitat may differ is in the prey selected from
the habitat.
Fourth, structural habitat as determined by first sightings is a
static concept: it says nothing about how an animal gets from one
part of its home range to another, nor indeed about how perches
of various heights and diameters are actually connected together
in the habitat. Knowledge of the movements of animals is essen-
tial for certain kinds of studies, for example, those on foraging
strategies, and may provide some indication of the type of prey
captured. This limitation for Jamaican Anolis is important: Trivers
(personal communication) has shown that valencienni differs
strikingly from some other Jamaican anoles (e.g., lineatopus) in
its active but cautious manner of searching for prey. The limitation
should be kept in mind when valencienni is compared to the other
species below.
Despite these disadvantages, the structural habitat has proven a
useful "summary statistic" in describing Anolis communities. It
has the great advantage that large populations of animals can be
sampled quickly, thus permitting several such populations to be
studied in a season. More balanced and detailed intralocality
studies of Anolis populations are better for many purposes but
are impractical for the study of the geographic variation of the
niche. Hopefully, as greater numbers of these detailed studies
4 BREVIORA JNO. 368
become available, it will be possible to set up correspondences —
e.g., animals which are often found on leaves take large numbers
of aphids — between the many properties measured in the detailed
studies and the few properties of the overall structural habitat.
Then some supposition can be made concerning the geographic
distribution of many more species characteristics than those ex-
plicitly given as the "structural habitat."
FORMAT
The format for reporting the results in this and succeedmg papers
will begin with a description of the localities studied, including a
list of the Anolis lizards seen. This will be followed by a verbal
summary of the results for each locality, accompanied by tables
depicting the structural habitat of the lizard classes considered, as
well as by a table indicating the statistical significance of differ-
ences in the habitat variables between all possible pairs of the
lizard classes. Finally, some relation will be made of the results
to previous studies of the species in question and to studies of the
habitats of species from other areas.
METHODS
Structural habitats were estimated by the "censusing" method
first developed by Rand (1964). In a transect through the study
area, each new lizard seen, unless obviously disturbed by the
observers, is noted as to its perch height, diameter, and (some-
times) insolation. Rand (1964, 1967a) and Schoener (1968)
have pointed out the possible errors in this technique, including
the especially serious one of differential visibility of various por-
tions of the habitat. Thus lizards, sitting, for instance, on the tops
of leaves in the canopy, are often likely to escape detection. How-
ever, the direction of difference between lizard groups within a
given site should not usually be affected by this kind of error. We
did most of the observations as a pair, using binoculars, the two of
us crisscrossing the habitat about 10 to 20 feet apart; this technique
probably reduced considerably the chance of missing lizards. Study
areas were often censused several times per day, but as a rule no
part of the area was censused at intervals closer than one and a
half hours; observation of lizard behavior in the interval indicates
that this was ample time for the animals to "recover" from dis-
turbance.
1971 JAMAICAN ANGLES 5
Data were cast into standard tables for easy intergroup compari-
son of structural habitat (Tables 1-4). For each lizard class for
which microclimatic information was taken, observations of cli-
matic categories — sun, shade, and clouds — were lumped into a
single structural habitat table. Additionally, however, the percent
occurrence in the three climatic categories were Hsted separately
for each class (Table 5).
A powerful new technique of multivariate analysis was used to
compute the statistical significance of habitat differences among
the various groups of lizards. The technique utilizes the iterative
procedure of Deming and Stephan (1940) and was recently ex-
pounded by Bishop (1969). It is designed to detect associations
between variables — in our case perch diameter, perch height, lizard
class, and insolation — of complex contingency tables; thus it can
handle both nominal and ordinal variables. Because it also simul-
taneously considers associations between the habitat variables per
se, this technique should largely eliminate any apparent difference
in habitat among lizard classes caused entirely by the structure of
the vegetation. For example, were relatively high perches always
also relatively thin perches, a lizard class which occupied signifi-
cantly higher perches, in the statistical sense, would not necessarily
occupy significantly thinner perches, because of the strong height
versus diameter interaction. Details of the application of the
method to our data are given in the appendix (see also Schoener,
1970). The reader not famihar with these techniques should first
consult the paper by Fienberg (1970), which is an exposition of
the method written especially for ecologists.
THE SPECIES
There are seven species of Anolis described for Jamaica (Under-
wood and Williams, 1959).
Two of the species — sagrei and grahami — can be characterized
as inhabiting strictly lower and middle elevations. A. grahami is
very widespread, occurring abundantly throughout the lowlands.
Ty^icdX grahami (subspecies grahami) are medium-sized {' snout-
vent length (SVL) = 65.5 mm; 9 SVL = 44.0 mmy green to
1 Means are of the largest third of all specimens examined (see Schoener,
1969).
6 BREVIORA No. 368
green-blue lizards occurring in all areas but the northeast, where
they are replaced by grahami aquarum, a somewhat smaller
(^ SVL = 61.8 mm; 9 SVL = 45.1 mm), bright emerald green
lizard. A. sagrei, a medium-small (^ SVL = 50.4 mm; 9 SVL =
40.9 mm) brown species, is restricted in habitat on Jamaica, occur-
ring only over the western portion of the island and there confined
to the openest, sunniest areas. It is often associated with rocks.
Apparently, sagrei has invaded Jamaica from Cuba and is in the
process of spreading eastward (Underwood and Williams, 1959;
Williams, 1970).
Another primarily low and middle elevation species is lineatopus.
This medium-sized (^ SVL = 62.7 mm; 9 SVL = 43.6 mm)
lizard is perhaps the most varied in its coloration and pattern: four
subspecies are recognized, some of which are found in dry open
areas and others of which are restricted to the darkest forest
(Underwood and WilUams, 1959).
Two species have been recorded from the lowlands to c. 4500
feet. A. opalinus, slightly smaller than sagrei (^ SVL = 49.5 mm;
9 SVL = 40.5 mm), comes in varying patterns and shades of
brown and grey. There is scarcely a locality in Jamaica that does
not have this species, though in the lowlands it is restricted to the
shadiest areas and in the uplands is found in very open situations
(Underwood and WilUams, 1959; Rand, 1967; this paper). The
grey-white, medium-large (^ SVL = 79.4 mm; SVL = 68.5 mm) 9
valencienni seems not to reach the density that the aforementioned
species sometimes do but is found throughout Jamaica, including
some areas above 4000 feet. It appears to be commoner in open
than in heavily shaded situations, but by no means is absent from
the latter (see below).
The above five species are the ones which occurred commonly at
one or more of the three localities studied, and which are therefore
included in the comparisons to follow. A sixth species, the "green
lizard" garmani, was seen at each of the three study sites, but rarely.
It is the largest of the Jamaican anoles (^ SVL = 110.0 mm; 9
SVL = 82.5 mm) and occurs throughout the island at all eleva-
tions (Underwood and Williams, 1959). However, it is most
abundant relative to other Anolis species at middle elevations
(e.g., Trivers, MS, and below), though it is also known to be
common at certain lowland localities, such as the "ironshore" vege-
tation along the northwest coast. The seventh species, reconditus,
1971 JAMAICAN ANGLES 7
is restricted to middle and upland elevations and has only recently
been discovered (Underwood and Williams, 1959; Lazell, 1966).
LOCALITIES
Three lowland localities were selected for study, representing a
wet, a rather dry, and a mesic area.
The first, about ten acres in extent, was located east of the town
of Port Antonio, on Jamaica's northeast coast. Specifically, it ex-
tended over the northwestern edge of a point of land supporting
the ruins of an estate locally referred to as the "Folly." The vege-
tation was quite secondary: planted trees and shrubs, such as Ficus,
limes, palms, and mango trees intermingled with native vegetation
such as Tenninalia. This locality had, however, become consider-
ably overgrown, and there then existed, side-by-side, an area of
almost continuous canopy and oftentimes sparse understory on the
one hand, and an area of widely spaced trees and shrubs with
much low, tangled herbaceous vegetation on the other. Both areas
were studied and will be discussed separately as "Port Antonio
Open" and "Port Antonio Closed."
The locality was worked 30 June-1 1 July 1967. Despite the fact
that Jamaica was then undergoing one of its most intense droughts,
rain fell fairly frequently though not protractedly; the area averages
131 inches of rain per annum (Handbook of Jamaica, 1966). The
anoline species grahami aquarum, lineatopus lineatopus (or inter-
mediates betwen lineatopus and ahenobarbus) and, to a lesser
degree, opalimis and valencienni were abundant in the open area;
the species opalimis and lineatopus, and to a much lesser degree
valencienni, were common in the closed area.
The second study site, about four acres in extent, was located at
Mona, near Kingston, in tall, open forest near the base of Long
Mountain. It apparently was marginal to Rand's (1967) "Mona
bush" study area, but differed in its much greater preponderance
of grahami. Vegetation in this area consisted of large trees and
smaller woody shrubs of typical tropical dry forest aspect: thorns,
flattened canopies and small, numerous leaves. The understory, in
addition to the shrubs, was mostly grass of about one to two feet
in height, but certain patches were practically cleared while others
had a more varied herbaceous vegetation. The latter appeared seri-
ously affected by the drought. The site in general falls within
Asprey and Robbins' (1953) "dry limestone scrub forest."
8 BREVIORA No. 368
The area was studied 13-18 July 1967. Rainfall was almost non-
existent during this period. The nearby Hope Gardens record 51
inches of rain per year (Handbook of Jamaica, 1966), though
because of the extreme microgeographic variation in climate found
on the Greater Antilles, this figure may not be the same as that
for the study site itself. The anoline species grahami grahami,
lineatopus lineatopus and opal'mus abounded in the area. In addi-
tion, valencienni was not uncommon.
The third study area was located about one mile west of White-
house, near the southwest coast. It consisted of groups of trees
and shrubs of mesic to xeric aspect, which remained after partial
conversion into pastureland. The portion of this area closest to the
coast was planted in limes and pimentos; further upland, patches of
native trees merged gradually into unbroken forest. In places, con-
siderable grassy and rocky areas showed effects of heavy grazing.
The vegetation is labelled by Asprey and Robbins (1953) "culti-
vated pasture or second growth scrub," bordered by remnants of
"dry limestone scrub forest," or by forest transitional between that
and "wet limestone forest." The total area considered encompassed
about 15 acres, though parts of it were not included in the censuses.
The area was studied 21-27 July 1967. Weather was regular and
cychcal during that time, sunny mornings giving way to overcast
or partly cloudy afternoons, accompanied sometimes by heavy
downpours. A nearby town (Bluefields) logs 91 inches of rain
per annum (Handbook of Jamaica, 1966). The form grahami
grahami occurred throughout the study area. The other two com-
mon species, sagrei and opalimis, were restricted to open and closed
areas respectively, and their horizontal ranges, though interdigitat-
ing, overlapped Uttle at the same time of day. A fourth species,
valenciemii, was seen rarely. The investigation of the opalimis-
grahami area has been partly reported in a different context
(Schoener, 1970) but will be reiterated in entirety below so as
to standardize its results.
RESULTS FOR LIZARD STRUCTURAL HABITATS
In the following discussion, all comparisons, unless stated other-
wise, are statistically significant (Tables 6-9) as judged by the
technique described in the appendix. Statements in the text to
follow, such as lizards of species A "occurred higher" or "were
1971 JAMAICAN ANGLES 9
higher" than those of species B, should be interpreted as descrip-
tive of the modal individual or the bulk of the population rather
than of all individuals in the species' population. Large lizards
could always be distinguished as adult males and will generally be
referred to hereafter simply as "males." Smaller lizards included
mostly adult or subadult females but also included some subadult
males; they were all lumped into the class "female-sized lizards"
because they could not usually be distinguished in the field. When
sufficiently abundant, the smallest lizards — "juveniles" — are con-
sidered separately.
Mona (Tables 1, 6). At Mona, ten classes of lizards in four species
were considered.
The highest lizard species observed was valencienni: males occurred
higher than any other class, and female-sized valencienni were higher than
all but grahami males. A. grahami was the next highest species: its males
occurred higher than all classes of lineatopus and opalinns. The males of
opalinus occurred higher than female-sized grahami and all lineatopus.
Female-sized grahami were higher than all lineatopus and female-sized
opalinus. Male lineatopus occurred higher than female-sized opalinus and
grahami juveniles. Female-sized opalinus occurred higher than grahami
juveniles and female-sized or juvenile lineatopus. Juveniles of grahami were
higher than female-sized and juvenile lineatopus.
A. opalinus as a species was on perches of the greatest diameter; males of
this species were on significantly thicker perches than all classes but male
grahami, and only the latter occupied thicker perches than female-sized
opalinus. Male grahami were also on thicker diameters than any other
interspecific class. Female-sized opalinus and male lineatopus had about
the same perch diameters but were on thicker perches than valencienni
and female-sized or juvenile grahami and lineatopus. Female-sized grahami,
valencienni, and lineatopus all occurred on thin perches of nonsignificantly
different diameter. Juveniles of lineatopus and grahami occupied the small-
est perches, the latter the smallest of all.
Intraspecifically, in all four species males perched higher than did female-
sized individuals, and female-sized individuals in grahami and lineatopus
occurred higher than did juveniles. In all but valencienni, males frequented
thicker perches than did female-sized individuals, and in grahami and
lineatopus, female-sized lizards were on thicker perches than juveniles.
Although climatic observations were not recorded at Mona, our impres-
sion is that the species tended to separate the way Rand (1967a) has de-
scribed for a nearby area: grahami and valencienni were in the openest,
sunniest situations; lineatopus was intermediate; and opalinus was in the
shadiest areas. Relatively shady areas at Mona are associated with large-
diametered trees, and the comparatively small opalinus preferred trees
10 BREVioRA No. 368
whose diameters were bigger than those for any group but grahami males.
Port Antonio Open Area (Tables 2, 7). The classes studied in the open
segment of the Port Antonio site were the same as those observed at Mona.
In relative height, the lizard classes were arranged in nearly identical
fashion to those at Mona. Male valencienni were the highest, as before,
followed by grahami males, which were here significantly higher than female-
sized valencienni. Probably the greater number of low thin-branched shrubs
in relation to trees in the open area at Port Antonio is responsible for this
discrepancy with the Mona area. Next in height were male opaliniis, higher
than female-sized and juvenile grahami as well as all classes of lineatopus.
Female-sized grahami and opalinus were distributed over similar heights
and were higher than any class of lineatopus. Males of lineatopus were
higher than juvenile grahami, but the latter were higher than female-sized
or juvenile lineatopus.
In diameter, there was a major reversal from the pattern at Mona. Adult
male valencienni, the largest of the four species, occurred on diameters not
significantly thinner than those of grahami or opalinus males and signifi-
cantly thicker than those of male lineatopus. Males of opalinus at Port
Antonio were on thicker diameters than grahami males, and the latter
showed no significant difference from female-sized opalinus. Female-sized
opalinus were on thicker perches than female-sized or juvenile grahami,
lineatopus, or valencienni. Males of lineatopus occurred on larger-diam-
etered perches than did female-sized and juvenile grahami or female-sized
valencienni. Female-sized grahami occupied thicker perches than did female-
sized valencienni or female-sized and juvenile lineatopus. Female-sized
valencienni occurred on thicker perches than did female-sized lineatopus or
juveniles of grahami and lineatopus. Finally, female-sized lineatopus were
on thicker perches than grahami juveniles.
Within the same species, males of grahami, opalinus, and lineatopus were
found higher than smaller-sized individuals. In addition, female-sized anoles
were higher than juveniles in grahami and lineatopus. For each species,
classes whose individuals were of the largest size were found on thicker
perches than all classes of smaller individuals.
Once again, opalinus and grahami segregated by shade and sun, respec-
tively. And again, opalinus, a small species, occurred on bigger trunks and
branches than did the larger species grahami or lineatopus. A. opalinus was
also found close to the ground in shrubby, more tightly packed vegetation,
where it was able to perch in the shade. In such areas, which were scat-
tered in patches throughout the study site, grahami ranged higher, being
found on leaves and more exposed branches. As at Mona, male lineatopus,
similar in size to male grahami, were more often found on thinner perches.
Unlike the Mona situation where leaves were mostly small, the Port
Antonio site contained many broad-leaved herbaceous plants. In such vege-
tation, particularly on leaves, it was common to see female and juvenile
1971 JAMAICAN ANGLES U
grahomi (Table 2). It is possible that aquarum, which is that form of
grahaini inhabiting the wettest lowland areas (including Port Antonio), is
both smaller and of a brighter, more leafy green color because of its oppor-
tunity for occupying the more luxuriant green vegetation resulting from the
heavy rainfall.
Port Antonio Closed Area (Tables 3, 8). In this more shaded site, so few
valencienni were seen that they are all lumped into one category. Repre-
sentatives of grahami were also very uncommon and are therefore not
included. In addition, because of the much greater abundance of opalinus,
female-sized individuals are treated separately from juveniles in that species.
In height, differences between all possible pairs of lizard classes were
significant. The order of groups, from highest to lowest, was valencienni,
opalinus males, lineatopiis males, female-sized opalinus, opalinus juveniles,
female-sized lineatopus, and lineatopus juveniles.
In diameter, opalinus males were on thicker perches than any other group.
Female-sized opalinus, male lineatopus, and valencienni occurred on perches
not significantly different in diameters, and all were on thicker perches than
female-sized lineatopus or juveniles of opalinus and lineatopus. Female-
sized lineatopus occurred on thicker perches than did juveniles of opalinus.
Intraspecific relations paralleled those for the other two areas: in opalinus
and lineatopus, the larger the size of the lizard, the higher it occurred, and
the thicker were its perches.
Whitehouse (Tables 4, 9). The study site near Whitehouse is identical in
species composition to those near Mona and Port Antonio except that sagrei
replaces lineatopus. The only form of lineatopus in the vicinity of the site
is neckeri, an animal which seeks darker forest than opalinus. As stated
above, climatic observations were recorded at Whitehouse in addition to
those on structural habitat. Although reported in part elsewhere (Schoener,
1970), data treatment here differs in two major respects: the category
"clouds" was added to those of "sun" and "shade" for the climatic variable,
and "time" is not considered as an additional variable.
In this area again, grahami males and valencienni (all classes combined)
were found at the greatest heights. They were followed by female-sized
grahami, which occurred higher than any class of sagrei or opalinus.
Juvenile grahami were found higher than sagrei and male opalinus. Both
classes of opalinus perched at greater heights than did any class of sagrei.
Male grahami, male opalinus, and female-sized opalinus did not show
significant differences in perch diameter, though the first were on the thickest
perches. Male grahami were found on thicker diameters than male sagrei,
but the latter did not differ significantly from either class of opalinus. Male
sagrei occurred on thicker perches than did female-sized grahami, juvenile
grahami and valencienni. Female-sized sagrei, juvenile sagrei and valen-
cienni all had greater-diametered perches than did juvenile grahami. In
addition, sagrei juveniles perched at greater diameters than did the com-
bined valencienni.
12 BREVIORA No. 368
Within the same species, adult males of grahami, sagrei, and opoliniis
perched higher than female-sized individuals. The latter in sagrei perched
higher than juveniles, but in grahami the two classes were similar in height.
Once again, males of the three commonest species occurred on thicker
perches than did smaller individuals, but the results were not significant for
opalinus. Juveniles in grahami and sagrei were found on thinner perches
than the other intraspecific classes.
Although there was an overall tendency for sagrei and valencienni to be
found most often in the sun and opalinus most often in the shade or on
cloudy days, this pattern varied by species class (Table 5). Males of
opalinus were most consistently found in the shade: they were seen signi-
ficantly more often during cloudy days or occurred more often on shady
perches than male and juvenile sagrei, male and female-sized grahami, and
valencienni. Juveniles of sagrei, in contrast, were very frequently seen in
the sun: all classes of grahami and male opalinus were seen significantly
more often in the shade or during cloudy weather than were juvenile sagrei.
Only two other intraspecific comparisons were significant: female-sized
grahami were most often seen in the sun or during cloudy days than male
sagrei, and valencienni were most often seen in the sun or on cloudy days
than male grahami. Thus there was no invariant tendency for that class
recorded most often in the shade to be also the one recorded most often
during cloudy weather.
Only a few intraspecific differences were significant: male opalinus were
recorded more frequently in the shade or on cloudy days than smaller
opalinus; female-sized grahami were more frequently seen in the sun or on
cloudy days than male grahami, and female-sized sagrei were more fre-
quently seen in the shade or during cloudy weather than juvenile sagrei.
Thus there was some inclination for the smaller-sized lizards within a species
to be on sunnier perches.
DIFFERENCES BETWEEN HABITAT VARIABLES
The statistical treatment of the structural-habitat data also de-
tects significant interactions between the two perch variables, height
and diameter, and when available, the climatic variable. Thus it
answers the following kind of question: is there a significant asso-
ciation between thin perches and high perches for all perches
combined of the two lizard classes being compared?
At Mona, for most comparisons among lizard classes, small
perches occurred at greater heights than did large perches (Table
6). However, in the three of nine cases in which a significant
height-diameter interaction was found (male grahami with female-
sized grahami, male valenciemii, or female-sized valenciemu) , the
1971 JAMAICAN ANGLES 13
reverse was true. In the Port Antonio open area, on the other
hand, where trees were more widely scattered and there was much
low second growth, all significant associations were of small-
diametered perches with low heights. Whitehouse resembled the
Mona area more than Port Antonio in its relation of perch height
to perch diameter; low perches tended to be of larger diameter.
This interaction again reflects the vegetation structure: at White-
house, there were few tall trees and little herbaceous understory —
most large perches were therefore low, including the fenceposts
especially preferred by sagrei. The only statistically significant ex-
ceptions to this pattern were for sagrei juveniles, the most terrestrial
class of lizards on the site.
At Whitehouse, it was also possible to look for associations
between the climatic categories and those of perch height and
diameter (Table 9).
There were few significant interactions between perch height and
insolation, probably because, in the patchy vegetation of the study
site, the sun penetrated for the most part to vegetation of all
heights. For intraspecific sagrei comparisons, higher perches
tended to be relatively shady and lower perches relatively sunny.
Individuals of sagrei were found more often than those of other
species in areas away from the shade provided by large trees and
shrubs: in such areas, most perches are both low and sunny. Lower
perches in sun than during cloudy weather were found for the com-
bined data of juvenile grahami and male sagrei. In contrast, the
lowest perches were found in the shade or sun and the highest on
cloudy days for female-sized sagrei lumped with grahami males.
Why these were the only interspecific comparisons showing a sig-
nificant height-insolation interaction is not apparent.
There were many more significant associations between perch
diameter and insolation. All but one were of two sorts: the thinnest
perches were occupied in the sun and the thickest in either shade or
on cloudy days. (The exception was for female-sized and juvenile
grahami: thinnest perches were shadiest, and thickest perches were
utilized during cloudy weather.) In a previous study which also
separated observations by time of day, a three-way interaction be-
tween diameter, insolation, and time was detected at Whitehouse
(Schoener, 1970). Exterior perches tend to be thinner in all habi-
tats, but in patchy ones such as that at Whitehouse, tend to be
sunnier as well — thus the association may simply reflect the phys-
iognomy of the vegetation. However, Jenssen (1970) found that
14 BREVIORA No. 368
individuals of Anolis nebiilosus climb into vegetation during mid-
day and has attributed this behavior to a warming of the substrate.
Perhaps a similar thermoregulatory function can explain the lizards'
avoidance of sunny, large surfaces during most of the day at
Whitehouse.
In several of the comparisons for each locality, the statistical
procedure indicated that there might be significant three-way inter-
actions between the variables (see appendix). Several are of inter-
est. Two were interactions between perch diameter, insolation and
lizard class. These were cases in which a low class, one of sagrei,
was paired with a higher class, one of either grahami or valencienni.
In sagrei, thick perches are more likely to be used in the shade on
sunny days, whereas in the more arboreal forms, thick perches arc
more likely to be used on cloudy days. Another three-way inter-
action showed that sagrei males tended to seek out shady perches
that were most often relatively high, whereas female-sized sagrei
found their shady perches relatively lower. Both these results
probably reflect differences in regard to relative availability of sun-
less perches between the habitats of the classes being compared.
As inspection of Tables 6-9 shows, there is a considerable num-
ber of significant interactions involving cHmatic and/or habitat
variables alone; therefore the extended statistical treatment given
the data of this paper is well justified. Most of these interactions
reflect differences in the vegetational structure and its exposure to
the sun. As mentioned, a few may indicate thermoregulatory be-
havior on the part of the hzards. However, the use of the climatic
categories — sun, shade, and clouds — can only give a crude first
approximation of the climatic preferences of these animals. Finer
resolution would be gained were temperature, humidity, wind
speed, and other variables measured at each perch.
DISCUSSION OF LIZARD STRUCTURAL HABITATS AND
RELATION TO PREVIOUS STUDIES
The climatic and structural habitats of the lowland species can
be summarized as follows.
1. Climatic. As first pointed out by Rand (1967a) for Kings-
ton populations, in all localities grahami inhabited relatively open,
sunny places and opalimis relatively closed, shaded places. At
Mona and Port Antonio, Uneatopus was intermediate in this regard.
However, sagrei, its structural habitat counterpart at Whitehouse,
occurred in sunnier, more exposed places than did grahami. In that
1971 JAMAICAN ANGLES 15
locality, opaliniis occurred on lower perches than elsewhere and
thus occupied in part perches where Uiieatopus would have been
expected were it present.
2. Structural. In all four study areas, valencienni as a species
was seen higher than its congeners (Figs. 1-4). Wherever found,
grahami was next in height, followed usually quite closely by opa-
linus. However, at Port Antonio female-sized lizards of grahami
aquarum were lower than either class of opalinus, reflecting their
abundance in herbaceous vegetation at that site. Lineatopus and
sagrei both occurred lowest in their respective sites though sagrei
appeared relatively more terrestrial. The ranking of species by
height at Mona was the same as that found by Rand in several
localities near Kingston. As can be seen from the figures, no par-
ticular between-species relationship of species-size to height was
evident. However, within species, smaller lizards always tended to
occur lower than did larger ones.
The ordering of species with respect to perch diameter is slightly
less consistent from locality to locality. At Mona, both male and
female-sized lizards taken separately showed a perfect inverse rela-
tion of body size and perch diameter: the largest species, valen-
cienni, occurred on the thinnest perches, followed by lineatopus,
then grahami, and finally, the smallest species, opalinus (Fig. 1,
Table 1) . At Port Antonio Open, the situation was the same ex-
cept that male valencienni occurred on thicker perches than all but
male opalinus, and female-sized lineatopus occurred on thinner
perches than female-sized valencienni (Fig. 2, Table 2). At Port
Antonio Closed, where only two species were considered in detail,
opalinus again took thicker perches than did lineatopus for both
sexes (Fig. 3, Table 3). At Whitehouse, the ordering was essen-
tially the same as at Mona, except that sagrei replaces lineatopus
(Fig. 4, Table 4). However, sagrei is smaller than grahami, so
there is not a perfect inverse relation of species-size and diameter
at Whitehouse. Taken as a whole, the data are in almost total
opposition to what would be expected if perch diameters were
selected by species on the basis of body weight.^ The inverse rela-
tion is made all the more interesting by the fact that within each
1 It should be pointed out, however, that garmani, the largest species on
Jamaica, was very rare and was therefore not considered in the study sites.
What few data exist on the perch diameter of this species (Rand, 1967a,
our unpublished data) indicate that garmani do not inhabit thinner perches
than valencienni but rather are often found on large trees and therefore
often perch on large branches.
16 BREVIORA No. 368
species, all significant associations are of larger individuals with
larger perches and vice-versa. Clearly some explanation other than
a purely supportative one must be sought for species-specific differ-
ences in perch diameter.
A possible explanation is the following. Given that, within
species, larger individuals are found on larger perches, an inverse
relation between perch diameter and species size would be expected
if species evolved so that that class of a given species overlapping
in space the most with a class of another species was the one which
differed the most in size from the latter class. In other words, a
direct relationship between size and perch diameter within species
and an inverse one between species is one way of ensuring that
interspecific spatial overlap is between dissimilarly sized individuals.
In fact, given the direct, within-species relationship, none of the
24 possible permutations of the species ordered by diameter results
in less total difference in the sizes of the most closely overlapping
interspecific pairs, though a few alternatives are about as good as
the one discussed. Because anoline lizards of different sizes take
differently sized foods (Rand, 1967a; Schoener, 1967, 1968;
Schoener and Gorman, 1968), such staggering of sizes in space
should alleviate resource competition (Rand, 1967b; Schoener,
1968).
Reversal of both the within- and between-species relationships is,
of course, an alternative way of juxtaposing dissimilarly sized
classes from different species. Why then does this second arrange-
ment not occur instead? An answer can perhaps be found if we
examine the probable course of faunal increase on Jamaica. It is
highly unlikely that the four species evolved simultaneously and
sympatrically. Therefore, while in isolation from other anoles, the
first species to have evolved probably showed a direct within-species
relationship between body size and perch diameter: not only is a
direct relationship adaptive over the large range in body sizes
spanned by the different age classes, but dominant individuals
in Anolis are usually largest (Rand, 1967b; Trivers, in prep.;
Schoener, in prep.) and would therefore appropriate the most suit-
able perches. Furthermore, all solitary species studied in hetero-
geneous vegetation showed such a direct relationship (Rand and
Rand, 1966; Schoener, 1967; Schoener and Schoener, in prep.).
Upon coming together, in order to achieve the second arrangement,
the species would have to change their species-specific size and/or
1971 JAMAICAN ANGLES 17
perch diameters as well as the within-species relationship between
size and perch diameter. Rather than that, it seems more feasible
for species to shift their size and/or perch diameters in such a way
as to preserve the intraspecific relationship and still avoid overlap
of similarly sized individuals, i.e., in accordance with the existing
arrangement. The positioning of a relatively large species on
perches of relatively small diameter could then be facilitated by
morphological changes in body proportions, such as those in rela-
tive leg length. Indeed, some proportional differences do seem to
exist in the Jamaican species: valencienni, the largest species con-
sidered, has relatively short limbs, particularly in femur. Similar
changes in proportions would likely be more difficult to build into
the ontogeny of single species, as would be necessary were the
second arrangement adhered to.
Though there is no between-species relationship of perch height
and size, the within-species tendency for smaller individuals to be
found relatively low also results in minimal interspecific spatial
overlap of similarly sized individuals, except for the valencienni-
grahami combination, in which female-sized valencienni are about
the same size as male grahami. However, those classes occur on
quite different perch diameters (Figs. 1-2, Tables 1-4), and Trivers
(personal communication) has evidence for major differences in
searching for prey and, possibly, prey taxa between the two species.
Rand (1967a) noted a tendency for opalinus and lineatopus to
juxtapose dissimilar sizes according to height, in localities around
Kingston, but he found the opposite for grahami and lineatopus.
A further difference between Rand's and our study is that his data
for small-sized grahami are bimodally clustered by height, one
mode being below six feet and the other above ten feet. A likely
reason for the discrepancy is difference in the structure of the avail-
able vegetation of the respective sites. Rand observed most of his
grahami on the campus of the University of the West Indies where
there is little high, shrubby understory surrounding the large trees.
Thus the distribution of female-sized grahami may have paralleled
differences in vegetational layers. The three localities we looked
at were all more overgrown and vegetationally more heterogeneous.
We also found a greater difference in perch height between male
and female-sized grahami in all localities than did Rand. Possibly
at Mona there were many more females on high branches than we
were able to detect, but this is unlikely to be much of a factor at
18 BREVIORA No. 368
either Port Antonio or Whitehouse, where the canopy is lower and
more broicen. Again, the lack of a viney, bushy understory in the
"park-like" vegetation where Rand studied most of his grahami is
probably responsible for the difference. A third possible reason for
the difference is simply some effect associated with the smaller
sample size Rand used, either one purely of sampling error, or one
related to a smaller range of times of day or weather conditions
than covered in our study.
ACKNOWLEDGMENTS
We thank T. A. Jenssen, A. S. Rand, R. L. Trivers, and E. E.
Williams for critical comments on the manuscript and S. D. Fien-
berg for statistical advice. We also thank C. B. Lewis and T. Farr of
the Jamaica Institute and I. Goodbody of the University of the West
Indies for assistance in the field. Research was supported by NSF
grants GB 3167 to the Committee on Evolutionary Biology, Har-
vard University, and NSF grants GB 6944 and B 01 9801 X to E. E.
Williams.
APPENDIX I
Statistical Appendix
Data of the sort presented above are ideally treated in the form
of a multiway contingency table, because variables are both ordinal
(perch height and diameter) and nominal (lizard group, climatic
category). In the case of nominal variables, there is no problem
in selecting categories (referred to as "levels"): there are two
lizard groups and three climatic categories (see above). However,
for ordinal variables, a continuous set of quantitative data must be
broken at one or more places in order to form categories. Because
of the multiphcative increase in the total number of cells in a con-
tingency table with increasing number of categories for a single
variable, two categories each were chosen for perch height and
perch diameter. That is to say, all observations less than or equal
to some number were cast into one category, and all those greater
than that number were cast into the other. The point at which the
data were broken was chosen by computing that number which
gave the maximum difference in cumulative frequency between the
two distributions of observations belonging to the hzard groups
being compared. Hence this procedure was designed to detect
1971 JAMAICAN ANGLES 19
maximum differences in height and diameter taken separately be-
tween the lizard classes, although because of interactions between
variables, it will not necessarily produce a maximum difference in
a combined model. The critical values were, of course, usually
much removed from those required to give maximum interaction
between the environmental variables themselves (e.g., perch height
and insolation). The critical value so determined for height or
diameter was generally different in different comparisons, and
ranged from one-fourth inch to four inches for diameter and
eleven inches to ten feet for height. An alternative procedure
would have been to choose the same intervals for all comparisons,
but given the great variation in lizard habitat preference, would
have obscured most differences. What we have in effect done is to
redefine "high" and "thick" for each comparison. There are sta-
tistical objections to this procedure, but at present appropriate
alternative methods of grouping data are not available (Fienberg,
1970). The points at which the data were broken are fisted here
(Table 10) for two reasons: 1) Other researchers may wish to
define "high" and "thick" differently and therefore can better
compare their method with the one used here; and 2) It is of
biological interest when comparing habitat distributions to know
where the point of maximum difference fies, especially in case
the observations were repeated later in the same or similar areas.
Once categories were chosen, the procedures diverged for three
and four variable situations.
In the four-way case, a contingency model was first set up which
contained all possible two-factor or pairwise interactions between
the four variables; in this case there were ( ^ ) = 6 such inter-
actions. Then an iterative procedure described" elsewhere (Bishop,
1969; Mosteller, 1968; Fienberg, 1970; Schoener, 1970) was used
to fit the data to the model, that is, to compute expected values for
each ceU of the contingency table. Two measures of goodness of
fit, the standard chi-square and the log-likelihood chi-square (Kull-
back, 1959) were computed and degrees of freedom determined as
described by Ku and Kullback (1968) and Fienberg (1970). It
was then noted whether the model gave a fit satisfactory at the 5
per cent level. In most cases the two statistics were very similar,
but where they allowed a different conclusion to be drawn about
significance, the log-likelihood ratio chi-square was followed.
Next, each two-way interaction was individually dropped, giving
six new models. For each of these, the difference between the new
20 BREVIORA No. 368
model and the original model was evaluated for statistical signifi-
cance by testing the difference in their log-likelihood ratio chi-
squares, according to the partitioning technique expounded in
Kullback (1959), Ku and KuUback (1968), and Fienberg ( 1970).
If all new models were significantly diiferent from the old model
at the 5 per cent level the process was terminated. Otherwise, that
new model was then chosen (and thereby the corresponding inter-
action removed) whose log-likelihood ratio chi-square was most
similar to that of the original model. The procedure was then re-
peated, five new models each containing four two-way interactions
being tested against the model containing five two-way interactions.
Interactions were thus removed, one at a time, until all models
with a smaller number of interactions were judged significantly
different from the next most inclusive model, or until no interaction
remained.
Because of space limitations, detailed results could not be re-
ported as they were in a previous paper (Schoener, 1 970) . Instead,
the results are summarized in Table 10. The six tiers of the table
correspond to the six possible two-way interactions. For each lizard
combination, these are given a number from zero to four. A "1"
denotes that the interaction remained significant in the above sense
every time it was tested in the removal procedure. A "2" denotes
that the interaction was significant at least at the termination of the
procedure. A "3" means that the interaction was significant when
removed from the most inclusive model (with six interactions) but
not at termination. A "4" indicates that the interaction was sig-
nificant sometime during the procedure but not at the beginning or
end. A "0" indicates that the interaction was never significant.
Interestingly, in Table 1 0 most interactions could be labelled either
"0" or "1", and in the simpler three-variable case, all interactions
could be so labelled (Tables 7-9). In the discussion of the text,
any interaction labelled 1-4 is considered significantly non-zero,
but the tables should be checked for fine distinctions.
In the case of three-variable tables (perch height, perch diameter,
and lizard group), there are ( ^ ) =3 two-way interactions. The
reduction procedure for these was similar to that described for four-
way tables, but of course is much shorter: only three new models
need be tested against the most inclusive model on the first round
instead of six.
Rarely, a set of models was encountered which never gave a
1971 JAMAICAN ANGLES 21
chi-square value denoting a satisfactory fit of the model at the 5
per cent level, regardless of what interactions were removed. These
are labelled in the tables. In such cases, differences between models
were still computed in the usual way and the results listed in the
tables. In addition, however, a search was programmed for signi-
ficant three-way interactions, in order to see if an improved fit
could be obtained. In the case of four variables, there are ( 3 ) =
4 three-way interactions. The procedure was to test each of four
models corresponding to the addition of a different three-way inter-
action to the model with all two-way interactions. For the White-
house data, there was no tendency for any particular three-way
interaction to predominate: each produced the best fit at least once.
Some of these are discussed in the text. In no case was it necessary
to consider more than one three-way interaction in order to pro-
duce a satisfactory fit. Once such a fit was obtained, the three
two-way interactions able to be removed were deleted one at a
time as before, and differences in chi-square with more inclusive
models were tested. In most cases, the two-way interactions that
could be removed without producing a significant difference be-
tween models were the same as some of those removed in the
analysis of two-way interaction models only.
In the case of a model with three variables, there is but a single
three-way interaction. Fitting this interaction would be a trivial
exercise resulting in a perfect fit (within the hmits of computational
accuracy); therefore, three-way interactions could only be con-
sidered for tables with four variables.
In several cases it happened that margins (the total number of
observations in a particular category of a variable or combination
of such categories) were zero. For these cases, two procedures
were tried. The first was to correct for the additional degrees of
freedom lost in such a table according to the method of Bishop
(unpublished thesis; Fienberg, personal communication). Once this
was done, the removal procedure could be carried out as befora.
In no case in the present study was the recalculated number of
degrees of freedom zero or negative, though if there are too many
zeros this will happen. A second way to handle zero margins is to
adjust the table slightly by shifting one (if possible) or more ob-
servations so that margins are no longer zero. This is best done
conservatively, that is, so as to reduce the likelihood of achieving
a significant difference to the variables of interest, in our case those
22 BREVIORA No. 368
in structural habitat. For such tables, where there was a choice
from several cells for selecting the observation to be moved, the
cell with the most observations was chosen. While far less prefer-
able than the first method, table adjustment had to be carried out
for the three-variable case, because the initial 2X2X2 model
tested has, assuming no zero margins, but one degree of freedom;
thus no further deletion is possible. Unless otherwise specified,
values in the tables of significance for four-way comparisons are
computed by the first method, though in only about 15 per cent of
tables with zero margins so far examined did conclusions from the
two methods differ at all.
APPENDIX II
Remarks on Other Localities
Jamaica, like the other Greater Antilles, is large and topographi-
cally diverse, yet it contains only seven species as compared to
Puerto Rico's ten, Hispaniola's 24 and Cuba's 24. Jamaica's less
diverse fauna is in part apparently associated with a great variation
from locality to locality within Jamaica in regard to what species
are found in certain segments of the vegetation. Brief visits which
we made to other Jamaican localities give an inkling of this vari-
ability. Already shown is that in western Jamaica sagrei replaces
Uneatopus as the open-area trunk-ground Hzard. In darker, mesic
forests west of the Whitehouse locality (such as that near Ferris
Cross), the trunk-ground species is Uneatopus necked, an olive
green-brown form somewhat smaller than the nominate subspecies.
A. opalimis is also common in such forests, but inhabits less shady
places and is more likely to be encountered marginally. A . grahami
seems entirely absent from these dark forests, but garmani is pres-
ent. More xeric forests two to five miles east of the Whitehouse
study area contained no trunk-ground species per se. Instead,
opalinus occurred often on low perches (though no measurements
were made), and garmani seemed commoner than at any of our
study areas.
Yet other species combinations are possible. In natural beach
vegetation near Rose Hall on the northwest coast, we saw grahami
and sagrei commonly. Where this vegetation met the xeric "iron-
shore" limestone formation, these species were replaced by Uneato-
pus merope, a rubiginous form well camouflaged on the rust-colored
limestone, and garmani. In certain moist mid-elevation sites such as
1971 JAMAICAN ANGLES 23
can be found about Mandeville, garmani was the common arboreal
species, and opalinus occurred in quite open, sunny places, includ-
ing low woodpiles and fenceposts. In these localities we found
grahami to be practically absent, while lineatopus neckeri was com-
mon in the blackest part of the forest. E. E. Williams and T. A.
Jenssen (personal communication) have also observed opalinus in
exposed situations at Mandeville; Williams, however, ionnd grahami
moderately common in certain of these situations. In montane
forest (c. 4000 feet), such as that surrounding Green Hills, we
frequently observed opalinus in extremely exposed places, including
along roadsides as at least temporarily a terrestrial lizard. Other
lizards in the area were valencienni in open situations and garmani
in somewhat more enclosed places; we also saw several reconditus
in relatively dark woodland.
LITERATURE CITED
Andrews, R. M. 1971. The structural niche and Anolis polylepis. Ecol-
ogy, in press.
AsPREY, G. F., AND R. C. RoBBiNs. 1953. The vegetation of Jamaica. Ecol.
Monogr.. 23: 359-412.
Bishop, Y. M. M. 1969. Full contingency tables, logits, and split con-
tingency tables. Biometrics, 25: 383-400.
Deming, W. E., and F. F. Stephan. 1940. On a least squares adjustment
of a sampled frequency table when the expected marginal totals are
known. Ann. Math. Stat., 11: 427-444.
FiENBERG, S. E. 1970. The analysis of multidimensional contingency tables.
Ecology, 51: 419-433.
Jenssen, T. a. 1970. The ethoecology of Anolis nebulosits (Sauria, Iguani-
dae). J. Herpetology, 4: 1-38.
Ku, H. H., AND S. KuLLBACK. 1968. Interactions in multidimensional con-
tingency tables: an information theoretic approach. J. Res. Natl. Bur.
Standards— Math. Sci., 728: 159-199.
KuLLBACK, S. 1959. Information Theory and Statistics. New York, Dover
Publications. 399 pp.
Laska, a. L. 1970. The structural niche of Anolis scriptiis on Inagua.
Breviora, 349: 1-6.
Lazell, J. D. 1966. Studies on Anolis reconditus Underwood and Wil-
liams. Bull, Inst. Jamaica Science, Ser. 18: 5-15.
Mosteller, F. 1968. Association and estimation in contingency tables.
J. Amer. Stat. Assoc, 63: 1-28.
24 BREVioRA No. 368
Rand, A. S. 1964. Ecological distribution in anoline lizards of Puerto Rico.
Ecology, 45: 745-752.
. 1967a. The ecological distribution of the anoline lizards
around Kingston, Jamaica. Breviora, 272: 1-18.
. 1967b. Ecology and social organization in the iguanid lizard
Anolis lineatopus. Proc. U.S. Natl. Mus., 122: 1-79.
Rand, A. S., and P. J. Rand. 1966. Field notes on Anolis lineatus in
Curasao. Stud. Fauna Curasao, 24: 112-117.
SCHOENER, T. W. 1967. The ecological significance of sexual dimorphism
in size in the lizard Anolis conspersus. Science, 155: 474-477.
. 1968. The Anolis lizards of Bimini: resource partitioning in
a complex fauna. Ecology, 49: 704-726.
. 1969. Size patterns in West Indian Anolis lizards. I. Size and
species diversity. Syst. Zool., 18: 386-401.
. 1970. Nonsynchronous spatial overlap of lizards in patchy
habitats. Ecology, 51: 408-418.
Schoener, T. W., and G. C. Gorman. 1968. Some niche differences among
three species of Lesser Antillean anoles. Ecology, 49: 819-830.
Underwood, G.. and E. E. Williams. 1959. The anoline lizards of Jamaica.
Bull. Inst. Jamaica, Science Ser., 9: 1-48.
Williams, E. E. 1970. The ecology of colonization as seen in the zoo-
geography of anoline lizards on small islands. Quart. Rev. Biol., 44:
345-389.
1971
JAMAICAN ANGLES
25
HIGH
T
H
I
C
K
T
H
I
N
^Ct] j
o
LOW
©
Figure 1. Mona. Species-classes are ranked by perch height and perch
diameter; distances between species-classes are not representative of the
magnitude of difference. Circles have diameters in proportion to the length
of the individuals in the designated class. Clear circles are of classes found
mostly in open, sunny areas; shaded circles are of classes in mostly closed,
shady situations; intermediate classes are represented by half-shaded circles.
V = valencienni, G = grahami, O = opalinus, L = lineatopus.
26
BREVIORA
No. 368
HIGH
^^P
T
H
I
C
K
■smOH
H
I
N
^^ 1
®
LOW
Figure 2. Port Antonio Open. Symbols as in Fig. 1.
1971
JAMAICAN ANGLES
27
HIGH
T
H
I
C
K
^^ ]
o
T
H
I
N
®
LOW
Figure 3. Port Antonio Closed. Symbols as in Fig. 1.
28
BREVIORA
HIGH
No. 368
H
K
^^^B
i
T
H
I
N
LOW
Figure 4. Whitehouse. Symbols as in Fig. 1. S = sagrei.
97;
JAMAICAN ANGLES
29
Table 1. Mona.
Percent observations in various structural habitat categories.
H = >20'; G = ground; R = rocks; N = sample size.
Diameter
Ht.^~~--^in.)
(feet)
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
N = 385
10.5-20
5-10
3-4 3/4
<3
Total
N = 246
10.5-20
5-10
3-4 3/4
<3
Total
N = 31
10.5-20
5-10
3-4 3/4
<3
Total
N = 214
10.5-20
5-10
3-4 3/4
<3
Total
male grahami
H
= 2.3
G
= 0
R = 0
3.6 8.1
6.6
3.8
0
3.6 13.8
21.4
9.5
0.3
3.6 5.5
4.6
3.5
0
3.6 2.6
3.3
2.5
0
12.4 30.0
35.9
19.3
0.3
female-
-sized
grab
ami
H
= 0 G =
= 0.3
R
0.7
2.7
2.9
2.6
0
0.3
3.4
14.9
28.2
1.0
0.3
2.7
6.0
11.5
0
2.7
4.1
7.2
8.4
0
22.1
48.9
16.2
10.7
= 0
8.9
47.9
20.5
22.4
4.1
13.0
30.9
50.6
1.0
juvenile £r
ah
ami
H =
0
G = 0
R = 3.2
0 0
0
0
0
0 0
0
25.8
0
0 0
4.8
37.1
3.2
0 0
6.5
19.4
0
0
25.8
45.1
25.9
0
0
11.3
male
opalinus
H = 0
1.9
4.2
2.3
4.2
21.0
21.3
2.3
7.0
6.3
1.9
5.6
3.3
10.3
37.8
33.2
82.3
; = 0
1.4
12.4
3.5
0.5
17.8
3.2
R = 0
0
0.5
0
0
0.5
9.8
59.4
19.1
11.3
30
BREVIORA
No. 368
Table 1. (cont'd)
-~-~...,^^ Diameter
Ht>-~~^in.)
(feet)^---^
>5
5-2 1/2
2 1/4-7/8
7/8-1/8
leaves
Total
N = 235
female-sized
opalinus H
= 0
G
= 0.9
^ = 0
10.5-20
0.4
2.1
1.3
0.9
0
4,7
5-10
2.6
10.2
18.1
14.3
0
45.2
3-4 3/4
1.7
4.7
10.9
7.5
0
24.8
<3
3.4
8.1
8.3
4.9
0
24.7
Total
8.1
25.1
38.6
27.6
0
N = 15
juvenile opalinus H = 0
G =
0
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
22.2
0
22.2
3-4 3/4
0
0
0
11.1
0
11.1
<3
0
11.1
22.2
33.3
0
66.6
Total
0
11.1
22.2
66.6
0
N - 838
male
lineatop
us H = 0.2
G =
0.
8 R = (
)
10.5-20
0.4
1.1
1.8
0.1
0
3.4
5-10
2.6
13.4
22.6
13.0
0
51.6
3-4 3/4
2.1
7.2
14.1
9.5
0.1
33.0
<3
1.7
2.5
3.6
3.0
0
10.8
Total
6.8
24.2
42.1
25.6
0.1
K = 436
femal
e-sized
lineatopus
H = 0
G = 4.4
R = 0
10.5-20
0
0
0
0
0
0
5-10
0.5
1.4
6.0
3.9
0.2
12.0
3-4 3/4
0.5
4.6
8.4
18.9
0
32.4
<3
1.4
8.3
16.3
24.8
0.7
51.5
Total
2.4
14.3
30.7
47.6
0.9
1971
JAMAICAN ANGLES
31
Table 1. (cont'd)
Diameter
Ht. -^in.)
(feet)
N = 24
10.5-20
5-10
3-4 3/4
<3
Total
N = 15
10.5-20
5-10
3-4 3/4
<3
Total
K = 16
10.5-20
5-10
3-4 3/4
<3
Total
N = 3
10.5-20
5-10
3-4 3/4
<3
Total
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
juvenile lineatopus H=0 G=12.5 R=0
0 0
0 0
0 4.2
0 8.3
0
0
0
2.1
0
0
8.8
56.3
0 12.5
male valencienni
0 6.7
0 6.7
0 0
0 0
2.1
64.6
H = 6.7 G = 0
6.7 13.3
6.7 33.3
0 13.3
0 6.7
66.6
6.3
25.1
12.6
juvenile valencienni
0 0 0
0 0 0
0 0 0
0 0 0
50.1
= 0 G
33.3
0
0
66.7
100.0
0
0
0
8.3
0
0
12.5
75.0
8.3
26.7
46.7
13.3
6.7
female-
-sized
val
encienni
H = 6
3
G = 0
0
6.3
6.3
12.5
0
0
12.5
6.3
6.3
0
6.3
6.3
0
12.5
0
0
0
0
18.8
0
25.1
25.1
25.1
18.8
0
33.3
0
0
66.7
32
BREVIORA
No. 368
Table 2. Port Antonio Open.
Percent observations in various structural habitat categories.
H = >20'; G = ground; R = rocks; N = sample size.
Diameter
Ht>^~..,,.^(in.)
(feet)^^^-,^
>5
5-2 1/2
2 1/4-7/8
7/8-1/8
leaves
Total
N = 146
male
grahami
H = 4.1
G
= 1.4
R = 0
10.5-20
2.7
6.9
2.7
0
0
12.3
5-10
5.5
7.5
12.3
11.0
0
36.3
3-4 3/4
5.5
3.4
1.7
9.9
0.7
21.2
<3
4.8
2.1
6.8
8.9
2.1
24.6
Total
18.5
19.9
23.6
29.8
2.7
N = 163
female-sized <
grahami
H
= 0.6
G = 2.5
R = 0
10.5-20
0.6
0.6
0.6
0
0
1.8
5-10
1.2
2.5
7.7
5.2
0
16.6
3-4 3/4
1.2
0
2.8
10.7
1.8
16.6
<3
3.7
2.5
4.6
34.7
16.6
62.0
Total
6.8
5.5
15.6
50.6
18.4
N = 75
juvenile grab
ami H =
0
G = 0
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
0
0
0
3-4 3/4
0
0
0
10.7
0
10.7
<3
0
0
1.3
57.3
30.7
89.4
Total
0
0
1.3
68.0
30.7
N = 17S
male
opalinus
H = 1.1
G = 0
R = 0
10.5-20
0.6
1.7
10.8
1.7
0
14.8
5-10
5.7
15.9
12,8
4.3
0
38.6
3-4 3/4
6,3
11.4
5.4
6.0
0
29.0
<3
4.6
7.4
0.6
3.4
0.6
16.5
Total
17,1
36.4
29.6
15.3
0.6
1971
JAMAICAN ANGLES
33
Table 2. (cont'd) .
-...,,^^ Diameter
HtT~~-^-^4in . )
(feet) ^^-^-^^
>5
5-2 1/2 2
1/4-7/8
7/8-1/8
leaves
Total
N = 136
female
-sized opa
linus H
= 0 G =
= 0 R =
0
10.5-20
0
1.5
0
0
0
1.5
5-10
3.7
5.2
14.3
4.8
0.7
28.7
3-4 3/4
3.7
2.9
5.5
7.0
0.7
19.9
<3
2.9
6.6
11.0
27.9
1.5
50.0
Total
10.3
16.2
30.9
39.7
3.0
N = 21
juveni
le opalinus H = 0
G = 0
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0
4.8
4.8
0
9.5
3-4 3/4
0
0
0
14.3
0
14.3
<3
0
0
9.5
66.7
0
76.2
Total
0
0
14.3
85.7
0
N = 183
male 1
ineatopus
H = 0
G = 3.3
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
1.1
1.6
3.8
0
6.6
3-4 3/4
2.2
6.6
5.2
13.4
0.6
27.9
<3
3.8
7.7
12.0
37.7
1.1
62.3
Total
6.0
15.3
18.9
54.9
1.6
N = 110
female
-sized lineatopus
H = 0
G = 11.8
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
0
0
0
3-4 3/4
0
0
0.5
5.0
0.9
6.4
<3
0
0.9
9.1
63.6
8.2
81.8
Total
0
C.9
9.5
68.6
9.1
34
BREVIORA
No. 368
Table 2. (cont'd)
Htr-~-~-^n.)
(feet) ^~~--.>.^
>5
5-2 1/2 2
1/4-7/8
7/8-1/8
leaves
Total
N = 23
juveni
le lineatopus
H
= 0 G =
26.1
R
= 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
0
0
0
3-4 3/4
0
0
0
0
0
0
<3
0
0
0
56.5
17.4
73.9
Total
0
0
0
56.5
17.4
N = 25
male valencienni
H =
4.
0 G = 0
R =
0
10.5-20
8.0
12.0
16.0
0
0
36.0
5-10
8.0
12.0
12.0
8.0
0
40.0
3-4 3/4
4.0
4.0
0
0
0
8.0
<3
0
0
4.0
8.0
0
12.0
Total
20.0
28.0
32.0
16.0
0
N = 36
female
-sized valencienni
H = 2.8
G =
0
R = C
10.5-20
2.8
0
0
0
0
2.8
5-10
2.8
2.8
8.3
19.5
0
33.4
3-4 3/4
2.8
8.3
0
11.1
2.8
25.0
<3
2.8
0
0
33.3
0
—
36.1
Total
11.1
11.1
8.3
63.9
2.8
N = 3
juveni
le valencienni
H
= 0 G =
0 R
=
0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
33.3
0
33.3
3-4 3/4
0
0
0
0
0
0
<3
0
0
0
66.7
0
—
66.7
Total
0
0
0
100.0
0
1971
JAMAICAN ANGLES
35
Table 3. Port Antonio Closed.
Percent observations in various structural ha±>itat categories.
H = >20'; G = ground; R = rocks; N = sample size.
Diameter
N = 185
male
opalinus
H = 1.6
G = 0.
5
R = 0
10.5-20
6.5
1.6
1.6
2.2
0
11.9
5-10
7.0
9.7
22.2
9.7
0
48.6
3-4 3/4
5.9
1.6
8.6
2.7
1.1
19.9
<3
1.6
2.7
9.2
3.2
0.5
17.2
Total
21.1
15.7
41.6
17.8
1.6
N = 206
fema]
.e-sized
opalinus
H = 1.0
G =
2.4
R = 0
10.5-20
0.5
0.5
0.7
1.2
0
2.9
5-10
8.3
3.4
9.5
15.8
0
37.0
3-4 3/4
5.3
1.5
5.8
6.8
0.5
19.9
<3
6.3
3.9
9.5
15.8
1.5
37.0
Total
20.4
9.2
25.5
39.6
1.9
N = 79
juvenile opal
inus H =
0 G =
0
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
1.3
1.3
19.0
0
21.5
3-4 3/4
0
0
0
17.7
0
17.7
<3
0
0
9.5
50.0
1.3
60.8
Total
0
1.3
10.8
86.7
1.3
N = 122
male
lineatopus H = 0
G = 0
R
= 0
10.5-20
0
0
0
0
0
0
5-10
4.1
3.3
15.6
14.8
0
37.7
3-4 3/4
5.7
3.3
10.2
18.4
0.8
38.5
<3
4.1
3.3
9.0
7.4
0
23.8
Total
13.9
9.9
34.8
40.6
0.8
36
BREVIORA
No. 368
Table 3 . (concl'd) .
"--■-^^Diameter
(feet) ^^^^
>5
5-2 1/2
2 1/4-7/8
7/8-1/8
lee
ives
3.6
Total
N = 112
female
-sized lineatopus
H = 0"
G =
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0.9
0.9
8.9
0
10.7
3-4 3/4
0
0.9
4.9
21.0
0.
9
27.7
<3
2.7
1.8
16.5
30.8
6.
3
58.0
Total
2.7
3.6
22,3
60.7
7
2
N = 32
juveni
le lineatopus
H =
0 G =
3.1
R
= 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
6.3
0
6.3
3-4 3/4
0
0
0
6.3
0
6.3
<3
0
3.1
9.4
53.1
18
8
84.4
Total
0
3.1
9.4
65.7
18
8
N = 13
valencienni
H = 0
G =
0 R =
0
10.5-20
7.7
7.7
15.4
7.7
0
38.5
5-10
0
0
15.4
15.4
0
30.8
3-4 3/4
15.4
7.7
0
7.7
0
30.8
<3
0
0
0
0
0
0
Total
23.1
15.4
30.8
30.8
0
197;
JAMAICAN ANGLES
37
Table 4. whitehouse.
Percent observations in various structural habitat categories.
H = 20'; G ■= ground; R = rocks; N = sample size.
Diameter
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
Ht.^'^-.^dn.)
(feet)
N = 219
10.5-20
5-10
3-4 3/4
<3
Total
N = 284
10.5-20
5-10
3-4 3/4
<3
Total
N = 54
10.5-20
5-10
3-4 3/4
<3
Total
N = 88
10.5-20
5-10
3-4 3/4
<3
Total
male
g
rahami
H
= 1.4
G
= 0.5
R = 0
1.8
1.8
2.3
1.4
0
2.7
9.6
21.5
9.1
2.3
0.5
7.3
10.1
4.1
0.5
1.8
10.1
7.1
4.3
0
6.8 28.8
41.0
18.9
2.8
fema]
.e-
-sized
2£
ahami
H
= 0
G = 2.5
R
0
0
0.4
0
7
0
0.7
2.8
14.1
21
8
4.6
1.8
1.8
5.8
7
9
2.5
1.4
9.9
12.3
7
7
1.1
7.3
45.2
22.5
28.3
= 0.4
1.1
44.0
19.8
32.4
2.9 14.5
32.6
38.1
8.2
juvenile gr
ahami
H =
0
G = 0
R = 0
0 0
0
0
0
0 0
3.7
38.9
0
0 1.9
1.9
31.5
0
0 0
6.5
13.9
1.9
0
42.6
35.3
22.3
1.9
12.1
J4.3
1.9
male
opalinus
H =
0
G =
= 0
R
= 0
0
0
0
0
0
1.1
6.8
18
2
4.6
0
2.3
13.6
15
9
1.1
0
1.1
20.5
9
1
4.6
1.
1
0
30.7
32.9
36.4
4.5 40.9
43.2
10.3
1.1
38
BREVIORA
No. 368
Table 4. (cont'd) .
^"^^Diameter
Ht7""^^^in.)
(feet)^<v^
>5
5-2 1/2
2 1/4-7/8
7/8-1/8
leaves
Total
N = 72
female
-sized
opalinus H
= 0
G
= 2.8
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
5.6
7.0
1.4
0
14.0
3-4 3/4
2.8
4.2
16.7
1.4
0
25.1
<3
2.8
31.9
19.5
4.2
0
58.4
Total
5.6
41.7
43.2
7.0
0
N = 10
juveni
le opal
inus H = 0
G =
0
R = 0
10.5-20
0
0
0
0
0
0
5-10
0
0
10.0
20.0
0
30.0
3-4 3/4
0
10.0
10.0
20.0
0
40.0
<3
0
10.0
10.0
10.0
0
30.0
Total
0
20.0
30.0
50. D
0
N = 263
male sagrei
H = 0 G =
6.5
R
= 2.7
10.5-20
0
0
0
0
0
0
5-10
0
0.8
1.5
0
0
2.3
3-4 3/4
0
6.5
8.4
2.3
0
17.2
<3
2.3
30.0
27.8
11.4
0
71.5
Total
2.3
37.3
37.7
13.7
0
N = 393
female
-sized sagrei H =
0 G
=
13.2 R
= 4.8
10.5-20
0
0
0
0
0
0
5-10
0
0
0.8
0.3
0
1.1
3-4 3/4
0
1.6
1.3
0.3
0
3.2
<3
2.5
31.8
22.5
21.0
0
77.8
Total
2.5
33.4
24.6
21.6
0
1971
JAMAICAN ANGLES
39
Table 4. (concl'd) .
Diameter
(feet)
N = 251
10.5-20
5-10
3-4 3/4
<3
Total
N = 7
10.5-20
5-10
3-4 3/4
<3
Total
N = 8
10.5-20
5-10
3-4 3/4
<3
Total
N = 7
10.5-20
5-10
3-4 3/4
<3
Total
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
juveni
le sag
rei
H
=
0
G = 31
9
R
= 4
0
0
0
0
0
0
0
0
8
1.6
0
0
0.8
0
4
2.0
0
2.0
17.8
12
6
25.6
0.
4
29.2
14.3 0 42.9 42.9
female-sized valencienni H = 0
0 0
0 0
0 0
0 0
0
0
37.5
0
0
25.0
0
37.5
37.5
62.5
0.4
G = 0
0
0
0
0
0
0
0
2.4
3.2
58.4
ma]
Le
va
lencienni
H =
0
G = 0
R =
0
0
0
0
0
0
14
3
0
28.6
42.9
0
0
0
14.3
0
0
0
0
0
0
0
juvenile valencienni
H
= 0
G =
= 0
R =
0 0
0
0
0
0 0
42.9
14
3
0
0 0
0
28
6
0
0 0
14.3
0
0
0
85.8
14.3
0
R = 0
0
25.0
37.5
37.5
0
0
57.2
28.6
14.3
57.2
42.9
40
BREVIORA
No. 368
Table 5. Percent observations for Whitehouse
species in climatic categories.
Sun
Shade
Clouds
male sagrei
17.4
74.4
8.3
female-sized sagrei
15.9
73.8
10.2
juvenile sagrei
33.8
58.8
7.4
male grahami
12.4
74.6
12.9
female-sized grahami
20.5
61.4
18.1
iuvenile grahami
15.7
66.7
17.6
male opalinus
3.5
72.1
24.4
small opalinus
14.5
66.3
19.3
valencienni
36.4
50.0
13.6
1971
JAMAICAN ANGLES
41
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JAMAICAN ANGLES
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No. 368
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JAMAICAN ANGLES
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No. 368
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BREVIORA
Mitasemiinii of Comparative Zoology
Cambridge, Mass. 29 January, 1971 Number 369
LITHOPHAGA ARISTATA IN THE SHELL-PLATES OF
CHITONS (MOLLUSCA)
Robert C. Bullock
and
Kenneth J. Boss
Abstract. The occurrence of the mytilid bivalve Lithophaga aristata as
a borer into the shell-plates of polyplacophorans is unreported. Our investi-
gation revealed this lithophage in the Panamic Chiton stokesii and, less
commonly, in the West Indian C. tiiheniihitiis. A review of other organ-
isms known to associate with chitons is provided, although none is appar-
ently deleterious to the host-species as in the case of Lithophaga. Damage
to the chiton by L. aristata appears to consist predominantly of the weaken-
ing of the shell-plates and not the destruction of the aesthetes of the
nervous system.
INTRODUCTION
Although several symbiotic organisms are known to live in
association with chitons, the boring of the mytilid bivalve Litho-
phaga into the shell-plates of polyplacophorans appears to be
unreported in the literature. During a dissection of a specimen of
Chiton stokesii Broderip, 1832, several Lithophaga {Myojorceps)
aristata (Dillwyn, 1817) were discovered in the shell-plates. A
survey was undertaken to assess the occurrence, and to determine
the nature, of the infestation of Lithophaga in C. stokesii and
other species of chitons from various localities. In addition, the
known symbionts of chitons were reviewed. The results, presented
in this paper, indicate that the presence of Lithophaga in the shell-
plates of chitons is unusual and represents the most detrimental
polyplacophoran symbiont known. We have observed L. aristata
boring in the shell-plates of the Panamic Chiton stokesii and the
Caribbean Chiton tuberculatiis Linnaeus, 1758.
BREVIORA No. 369
ACKNOWLEDGMENTS
The manuscript has been critically read by our colleagues in
the Museum of Comparative Zoology, Messrs. R. I. Johnson and
M. K. Jacobson, and Dr. Peter W. Glynn of the Smithsonian
Tropical Research Institute. The X-ray apparatus was made avail-
able by the Museum of Comparative Zoology through a Milton
Fund grant to Drs. R. D. Turner and S. J. Gould. Funds enabhng
the senior author to collect Panamanian Polyplacophora were
provided by a National Science Foundation grant, GB 8620, issued
to the Committee on Evolutionary Biology, Dr. R. Rollins prin-
cipal investigator.
METHODS AND MATERIALS
Numerous specimens of Chiton stokesii, collected at various
localities in the Gulf of Panama by the senior author during July
and August 1969, were examined by radiographic techniques for
the presence of Lithophaga. A survey was made of the large col-
lection of Polyplacophora in the Museum of Comparative Zoology,
Harvard University, in an effort to locate other species host-
ing Lithophaga, and X-rays of suspected host-individuals were
made. All radiographs were checked for the number of lithophages
present, their spatial distribution, and orientation. Borers were
extracted at random to check their identity and all proved to be
L. aristata.
POLYPLACOPHORAN SYMBIONTS
Few records are to be found in the literature concerning sym-
bionts of polyplacophorans, but organisms representing several
phyla are associated with chitons. Cryptochiton stelleri (Midden-
dorff, 1846) is known to harbor two commensals: a crustacean,
Opisthopus transversus (Rathbun, 1893) and an annelid, Arctonoe
vittata (Grube, 1855) (MacGinitie and MacGinitie, 1968; Web-
ster, 1968).
Glynn (1968) and Menzies and Glynn (1968) summarized
present knowledge of the symbionts of the mantle cavity on the
West Indian chitonids, Acanthopleura granulata (Gmelin, 1791)
and Chiton tuberculatus Linnaeus, 1758. Included were: the
isopods, Dynamenella perforata (Moore, 1901) with A. granulata
and C. tuberculatus; Exosphaeroma alba Menzies and Glynn,
1971 LITHOPHAGA IN CHITONS 3
1968, Exosphaeroma cremilatum (Richardson, 1902), Dynamen-
opsis dianae Menzies, 1962 with C tuberciilatus; Exosphaeroma
alba var. chromata Menzies and Glynn, 1968 with C. mannoratus
Gmelin, 1791; and the foraniiniferan, AcerviiUna inhaerens
Schulze, 1854 and the amphipod, Parhyale hawaiensis (Dana,
1853) both with C. tuberciilatus; the harpacticoids, Harpacticus
sp. and Heterolaophonte sp. with A. ^ranulata and C tuberciilatus;
and the coUembolan, Actaletes ueptiini Giard, 1889 with A.
gramilata.
The mite Halixodes chitonis (Brucker, 1897) has been found
attached to the gills of the Neozelanic Cryptoconchus porosus
(Burrow, 1815) (Brucker, 1897; Brucker and Trouessart, 1900).
Helfman (1968) observed the ctenostomatous ectoproct Farella
elongata (van Beneden, 1845) in the ventral girdle tissue border-
ing the pallia! groove. Arey and Crozier (1919) reported the
following symbionts on the shell-plates of C. tuberciilatus: the
barnacle, Tetraclita; the polychaetes, Spirorbis and Serpula; and
algae, including the "Enteromorphas." They mentioned that the
algae afforded protection for various young moUusks, nematodes,
archiannelids, and protozoans. None of these symbionts has been
shown to be harmful to the host.
After conducting extensive studies on Chiton tuberciilatus , Arey
and Crozier (1919: 171-172) remarked: "The general impres-
sion derived from the consideration of destructive agents in relation
to Chiton is that these mollusks are very efficiently protected. The
length of life which they seem to attain, the variety of habitats
which they frequent, and the character of their sensory responses,
which determine certain features of their life in their habitats,
afford important evidence to this effect."
While the above statement is generally true for C. tuberciilatus,
the Panamic C. stokesii appears far more vulnerable to attack by
destructive agents than C. tuberciilatus. We observed large indi-
viduals of C. stokesii from different localities that were not only
greatly eroded, but heavily fouled with calcareous algae, bryozoans,
and polychaete tubes. Most of the latter were heavily infested with
Lithophaga aristata, some of which had grown large enough to fall
out of their burrows, leaving greatly weakened shell-plates. The
cases of C. tuberciilatus from Trinidad (MCZ 31955) and Isla
Margarita (MCZ 273763), in which we found examples harboring
L. aristata, appear to be rare. Our examination of numerous other
4 BREVIORA No. 369
samples of C. tuberculatus produced no additional Lithophaga.
It is probable that differences in shell-structure account for the
greater penetrability and susceptibility for fouling and boring in
C stokesii.
RESULTS AND DISCUSSION
The mytilid bivalve Lithophaga {Myojorceps) aristata (Dillwyn,
1817) (Figs. 7 and 8) bores into calcareous substrates, including
the shells of large bivalves (e.g., Spondyhis, Chama, Ostrea) and
gastropods (e.g., Haliotis, Patella, Strombus, and Pleuroploca) .
The species occurs in warm temperate to tropical waters in the
eastern Pacific, western Atlantic, and eastern Atlantic regions and
is usually found in shallow water, although Soot-Ryen (1955)
reported a living specimen taken from 165 fathoms (Turner and
Boss, 1962).
In both C. stokesii and C. tuberculatus, Lithophaga aristata
was present only in large specimens. This relationship coincides
with the findings of Arey and Crozier (1919) and Crozier and
Arey (1920) who observed the presence of barnacles, polychaetes,
and algae only on larger C. tuberculatus. The erosion of the shell-
plates, which is brought about by physical agents of the environ-
ment and by organisms which live on the chiton's shell, appears
to be a prerequisite for the boring of Lithophaga. During settle-
ment the pediveligers of L. aristata evidently reject the uneroded
areas of the polyplacophoran shell and metamorphosis occurs on
the eroded substrate. We noted that most lithophages began
boring at the posterior edge of the intermediate valves, which was
normally eroded in large individuals. Large C. stokesii showing
little or no erosion had few, if any, L. aristata; the valves of those
chitons that were considerably eroded revealed Lithophaga boring
in them at various places, not just at the posterior edge. Chiton
viridis Spengler, 1797, a Caribbean species that normally lives
below the low-water mark, is rarely eroded and no Lithophaga were
found in the shell-plates of this species.
After initial penetration of the shell, most L. aristata bored hori-
zontally. In several instances we observed the burrows of Litho-
phaga extending into a second valve. In one example from Culebra
Island, Canal Zone, which harbored over 40 L. aristata (Fig. 1 ),
1971 LITHOPHAGA IN CHITONS 5
two individual lithophages had bored vertically from one valve to
the next, penetrating the intervening musculature (Fig. 4). One
such burrow passed from valve III into the insertion plate of valve
IV (Fig. 1). All cases of vertical burrows apparently occurred in
overcrowded conditions.
Most burrows of intermediate valves were roughly parallel with
the antero-posterior axis of the chiton with the siphons of the
Litlu)pluii>a pointed posteriorly, although they deviated somewhat
by orienting themselves along an axis extending from the zone of
erosion toward the nearest growth zone. The reason for this is
obvious. Should a lithophage burrow perpendicularly to the antero-
posterior axis and only in the region of the mucro, it would risk
having its surrounding substrate eroded away. By burrowing from
the zone of erosion toward a growth zone, the lithophage is assured
of an increasing substrate in which to bore. This explanation clar-
ifies the otherwise haphazard orientation of the burrows in the
posterior valve, where the oldest shell material is near the center,
not at the posterior edge (Fig. 2).
As the lithophage increases in size, it faces problems caused
by the restricted space in which it can grow. Although some
Lithophaga enlarge their burrows dorsally, most penetrate ventrally
and eventually reach the mantle of the chiton. When the latter
situation occurs, the chiton secretes a thin calcareous shield in an
effort to contain the intrusion of the lithophage. Sometimes more
than half of the ventral portion of the lithophage is situated below
the ventral shell-plate surface. In spite of the efforts of the Lith-
ophaga to increase in size and the chiton's effort to contain it, the
lithophage soon reaches a point where further growth is impossible.
Whether this represents a truly stenomorphic condition or not is
uncertain, because we do not know if the L. aristata we observed
ever reached sexual maturity. Lithophaga aristata is known to
attain a length of 52 mm (Turner and Boss, 1962); the largest
specimen extracted from a C. stokesii was 9.0 mm, while the aver-
age length was about 7.0 mm.
There is evidence that at least a few L. aristata outgrow their
restricted polyplacophoran substrate (Fig. 6). Some of the largest
burrows that we examined were exposed along their entire dorsal
surface, indicating that the lithophages may have fallen from their
6 BREVIORA No. 369
burrows. It is probable that specimens that outgrow the chiton
perish in the external environment.^
The effect of numerous Lithophaga in the shell-plates must be
detrimental to Polyplacophora. In addition to damaging the spe-
cialized portion of the nervous system that makes up the aesthetes
in the tegmentum, Lithophaga weakens the valves and, at times,
probably affects the maneuverability of the chiton. The seriousness
of damage to a large amount of the tegmental nervous tissue is
open to question and may depend upon the species involved. Aes-
thetes are photosensitive, being activated both by light of constant
intensity and by a decrease of light intensity (Arey and Crozier,
1919). Studies on the growth and behaviour of Chiton tubercu-
latus indicated that normal erosion of shell-plates and concomitant
loss of tegmental aesthete photosensitivity produced at least a
partial inversion in its phototropic response. While young C. tiiber-
ciilatiis are photonegative, older individuals may be irresponsive or
photopositive (Arey and Crozier, 1919; Crozier and Arey, 1920),
although Glynn (personal communication) found older C. tubercu-
latus from Puerto Rico also to be photonegative. It appears, then,
that Lithophaga mainly affects the durability of the shell, rendering
the chiton more susceptible to predation.
REFERENCES CITED
Arey, L. B., and W. J. Crozier. 1919. The sensory responses of Chiton.
J. Exper. Zoo!., 29: 157-260.
Brucker, a. 1897. Sur un noiivel Acarien marin. C. R. See. Biol., 49:
632-633.
Brucker. A., and E. Trouessart. 1900. Seconde note sur un Acarien
marin (Halacaride), parasite de VAcanthocliiton porosiis. C. R. Soc.
Biol.. 52: 107 109.
Crozier. W. J., and L. B. Arey. 1920. On the ethology of Cliiton tiiber-
ctilatiis. Proc. Natl. Acad. Sci.. 5: 496-498.
1 Although Hodgkin (1962) maintained L. pitimiila kelseyi Hertlein and
Strong, 1946 outside their burrows for over one year under laboratory con-
ditions. Otter (1937) was unsuccessful in his attempts to rear L. cumingi-
ana (Reeve, 1857) and L. teres (Philippi. 1846) that had been removed
from their burrows. It is doubtful if Lilliophaga could survive out of its
burrow under exposed conditions.
1971 LITHOPHAGA IN CHITONS 7
Glynn, P. W. 1968. Ecological studies on the associations of chitons in
Puerto Rico, with special reference to sphaeromid isopods. Bull. Mar.
Sci., 18: 572-626.
Helfman, E. S. 1968. A ctenostomatous ectoproct epizoic on the chiton
Ischnochiton mertensii. Veliger, 10: 290-291.
Hodgkin, N. M. 1962. Limestone boring by the mytilid Lithophaga.
Veliger, 4: 123-129, 3 pis.
MacGinitie, G. E., and N. MacGinitie. 1968. Notes on Cryptochiton
stelleri (Middendorff, 1846). Veliger, 11: 59-61, pi. 6.
Menzies, R. J., AND P. W. Glynn. 1968. The common marine isopod
Crustacea of Puerto Rico. A handbook for marine biologists. Studies
Fauna Curasao, 27: 1-133.
Otter, G. W. 1937. Rock-destroying organisms in relation to coral reefs.
British Mus. (Nat. Hist.) Great Barrier Expedition 1928-29, Sci.
Repts., 1: 323-352, 6 pis.
Soot-Ryen, T. 1955. A report on the family Mytilidae (Pelecypoda).
Allan Hancock Pacific Expeditions. 20: 1-154, 10 pis.
Turner, R. D., and K. J. Boss. 1962. The genus Lithophaga in the
western Atlantic. Johnsonia, 4: 81-116, pis. 57-75.
Webster, S. K. 1968. An investigation of the commensals of Crypto-
chiton stelleri (Middendorff, 1846) in the Monterey Peninsula area,
California. Veliger. 11: 121-125.
BREVIORA
No. 369
1971 LITHOPHAGA IN CHITONS
Figure 1. Radiograph of disarticulated shell-plates from a specimen
of Chiton stokesii from Culebra Island. Panama Bay, Canal Zone
(MCZ 277122), showing a high degree of infestation by Lithopliaga aris-
tata (0.84 x ).
Figure 2. Enlargement of posterior valve of Fig. 1 (3.2 X ).
Figure 3. Typical siphonal opening of L. aristata burrow in an uneroded
portion of a C. stokesii shell-plate (12 X )•
Figure 4. Intervening shell-plate musculature located between valves IV
and V in Fig. 1, showing damage resulting from penetration of Lithophaga
(9.2 X).
Figure 5. Example of L. aristata (MCZ 277123) in shell-plate cross
section of C. stokesii from Punta Mala, Panama (3.9 x )•
10
BREVIORA
No. 369
Figure 6. Exposed burrow of Lithophaga aristata in Chiton stokesii from
Panama City, Panama (MCZ 78821) (4.2 x ).
Figures 7-8. L. aristata extracted from burrow illustrated in Fig. 5
(MCZ 277123) (6.5 x ).
BREVIORA
Miaseuni of Comparative Zoology
Cambridge, Mass. 31 March, 1971 Number 370
ECOLOGICAL OBSERVATIONS ON A LITTLE KNOWN
SOUTH AMERICAN ANOLE: TROPIDODACTYLUS ONCA
James P. Collins^
Abstract. The little known anole Tropidodactylus onca on the island
of Margarita is typically restricted to belts of low xerophytic vegetation
adjacent to the open sandy area of natural beaches. Most animals are
found on the ground or in low bushes and occur up to a height of 30.0
cm. They are poor climbers and will occasionally escape into holes made
by ghost crabs.
INTRODUCTION
A brief visit (from July 8 to July 21, 1968) to the Fundacion
La Salle on Margarita Island, Venezuela, afforded me an oppor-
timity to collect and observe a little known anole, Tropidodactylus
onca. Margarita is a continental island approximately twenty-one
miles off the district of Sucre on the northern coast of Venezuela.
It is approximately two hundred air miles northeast of Caracas.
Tropidodactylus onca is a speciaUzed derivative of Anolis but
with keeled scales underneath the digits instead of the expanded
digital pads with adhesive lamellae so characteristic of the latter
primarily arboreal genus. The difference in morphology should
be reflected in ecology, but there have been no detailed reports
on the habits, habitat, or even color in Ufe of T. onca. This
paper attempts to remedy this gap in information.
Description and color in life. Tropidodactylus onca is a rela-
tively large anole. The largest of the specimens collected is 75
mm snout- vent length. The tail is round with no dorsal crest
and is approximately equal to the snout-vent length of the animal.
The most distinctive specialization of the anoles in general,
the dewlap, is extremely well developed in this genus. It has a
bright yellow ground color in which individual red scales are
embedded.
1 Museum of Zoology, University of Michigan, Ann Arbor. Mich.
BREVIORA
No. 370
The dorsal color of the lizard varies from a very light ashy-
gray to a dark gray-brown with a disruptive pattern of black
and white longitudinal markings. These vary from barely visible
to very prominent. The ventral surface of the lizard is white.
The species is not sexually dichromatic and is very difficult to
sex externally. The cryptic coloration of the species is perfect.
In the field the animals are indistinguishable from their sur-
roundings. In fact, an animal can usually be located only when
the collector inadvertently frightens it into movement.
Habits and habitat. Tropidodactylus onca is a beach anole.
Its distribution is typically restricted to the belt of low xerophy-
tic vegetation adjacent to the open sandy area of a natural beach.
The exact width of this belt and its continuity varies according
to the geological and ecological factors of the particular locality.
The animal was never observed in the open sandy areas devoid
of vegetation, and penetrates only slightly into the more land-
ward areas where the vegetation is higher than 0.75 to 1.00 m.
20
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10
64"
50
10
50'-
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LOS FRAILES
El Aguo
Punta Montadaro
Playa Guacuco
ISLA CUBAGUA
ISLA COCHE
PENINSULA DE ARAYA
(0'
SO
Figure 1. Margarita Island. Localities at which Tropidodactylus onca
was observed or collected are shown as solid circles.
1971 TROPIDODACTYLUS ONCA 3
All the specimens, with the exception of one taken at a height
of 1.25 m, were collected on the ground or up to 30.0 cm high
on low bushes, vines, etc.
Observation and collecting was restricted to six localities (Fig.
1), all on the eastern half of the island. Roze (1964), however,
has reported the lizard from the western half (Macanao). Of the
six localities in which the species was observed, five are on the
coast. The exception, El Aguila, is a town approximately 2.5
km from Punta de Piedras. Possible reasons for the animal's
appearance here will be discussed later. Outside of this single
exception, no specimens were ever observed in noncoastal por-
tions of the island and all were collected at sea level.
The vegetation of the zone in which Tropidodactylus is found
consists of only low ground growth. The sand-shrub communities
of the five coastal localities in which Tropidodactylus is found
each contain at least two and as many as four of the following
species of plant: Sesuvium portidacastrum, Philoxerus vermicu-
laris, Bat is maritima, Sporobolus (virginicus?), Ipomoea (pescap-
rae?), and Mallotonia gnaphalodes. Howard (1950) lists these
six species as pan-Caribbean in distribution. These are found in
association with less abundant and restricted species, the exact
taxa varying from locality to locality. The landward edge of this
coastal community typically contains representatives of the genera
Opuntia, Melocactus, Lemairocereus, Philoxocereus, and Proso-
phis. Tropidodactylus penetrates slightly into this zone.
The majority of the animals collected were taken in large
patches of Ipomoea found at two of the localities investigated (El
Agua and Playa Guacuco). The reptile was found both on the
plants and in the open space between them. If the patch was
dense enough, the animals were typically found with their head
on top of the leaves and their trunk and tail among the tangled
vines. Some were also found on an occasional outcropping of
rock or log. When frightened, the animal would either duck into
the mat of vegetation (if dense enough) or scamper across the
open sand until it reached a vine or series of vines which it would
then proceed to run clumsily over. Being nonwoody, the plants
do not give much support and the animal could easily be taken.
In the area just north of Punta Montadero where Mallotonia, a
woody-stem plant, is dominant, the animal's behavior was very
different. Here, when first sighted, the lizard was always on the
BREVIORA
No. 370
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1971 TROPIDODACTYLUS ONCA 5
ground. When pursued, the majority of animals observed would
merely run among the ground cover. A few specimens, however,
were observed to climb the Mallotonia, some to a height of 30.0
cm. Their climbing was clumsy and ineffective. The toe structure
of this genus is not well adapted for tree climbing. Unlike most
anoles, T. onca is not arboreal.
Another means of retreat should also be pointed out. At times,
a specimen, being pursued, would run into a large hole in the
sand opening into a tunnel. Ruthven (1922) also reports this
species as escaping into holes. It should be noted, however, that
these holes are resting places made by ghost crabs (Ocypode) and
are not dug by Tropidodactylus. It should also be noted that this
was a rather infrequent means of escape, used by the lizard only
when almost completely exhausted.
At each of the six localities, T. onca is found sympatric with
Cnemidophorus lemniscatus lemniscatus. In those localities (El
Agua and Punta los Cocos) where the landward border of the
coastal area is occupied by a semi-desertic community, the terri-
tory of Tropidodactylus partially overlaps that of Tropidurus tor-
quatus hispidus. The Tropidodactylus penetrate this zone for only
a very small distance. In some areas, two other organisms also
found sympatric with Tropidodactylus are the gecko Gonatodes
vitatus vitatus and the microteiid Gymnopthalmus laevicauda.
Roze (1964) has the following note concerning the diet of
Tropidodactylus: "The stomach contents examined in various
specimens of this species revealed the remains of grasshoppers
(Grillidae), Coleoptera, spiders, and various species of Diptera, as
well as the remains of other unidentifiable arthropods." Tropido-
dactylus then, hke most anoles, is insectivorous. Similarly, like
most anoles, the animal is diurnal in its activity. All but one of
the twenty-five specimens were captured during the day. The
single exception was collected alongside the road near the town
El Aguila approximately 2.5 km from Punta de Piedras. It was
on a branch of a low bush, Jatropha gossypiifolia, in the cleared
margin alongside the road. The animal was in typical anole
sleeping posture, snout toward the main stem, but with its eyes
open. Just prior to being seized, the animal moved its head but
did not attempt to flee. This single exception to the otherwise
complete coastal distribution of the animals on the island most
probably migrated to this inland area along the corridor of low
vegetation bordering either side of the roadway. This habitat is
ecologically similar to that of the coastal zone.
6 BREVIORA No. 370
ACKNOWLEDGMENTS
I am grateful to Dr. Janis Roze and Dr. Ernest E. Williams for
critically reading the manuscript and for their valuable sugges-
tions. This study was carried out at the Fundacion La Salle, Mar-
garita Island, Venezuela; I thank Hermano Gines for making the
facilities there available to me. Field expenses were partially met
by NSF-GY-4183, administered by Manhattan College, and NSF-
GB-6944 to Ernest E. Williams.
REFERENCES
Howard, R. A. 1950. Vegetation of the Bimini island group. Ecol. Mono.
20: 317-349.
Roze, J. A. 1964. La herpetologia de la Isia de Margarita, Venezuela.
Mem. Soc. Cien. Nat. La Salle, 24 (69): 209-241.
RuTHVEN, A. G. 1922. The amphibians and reptiles of the Sierra Nevada
de Santa Marta, Colombia. Miscellaneous PubUcation No. 8, Museum
of Zoology, University of Michigan.
BREVIORA
Miaseiuioi of Comparative Zoology
Cambridge, Mass. 31 March, 1971 Number 371
A NEW SPECIES OF BROMELIAD-INHABITING GALLIWASP
(SAURIA: ANGUIDAE) FROM JAMAICA
Albert Schwartz^
Abstract. A new species of anguid lizard, Diploglossus fowleri, is
described from two specimens collected from bromeliads at the northern
edge of Jamaica's Cockpit Country. The affinities of the new species are
with D. hewardi and D. diiqiiesneyi; both D. fowleri and D. duquesneyi
appear to be geographic or ecological isolates of the widespread D. hewardi.
The Antillean islands of Jamaica and Hispaniola have excep-
tionally large numbers of species of the anguid lizard genus
Diploglossus Wiegmann. The latter island has six extant species,
whereas Jamaica likewise had six species of which one {occiduus
Shaw) is presently considered extinct. Cousens (1956) summar-
ized the then-known Jamaican galliwasps and regarded cnisculus
Carman, harbour i Grant, hewardi Gray, and duquesneyi Grant
as valid species. Since that time, D. microblepharis Underwood
has been named from a single specimen from the northeastern
Jamaican coast. Cousens (1956), followmg Grant (1940b), sep-
arated the four forms then recognized into two major groups:
one group {crusculus, barbouri) with short legs and the other
{hewardi, duquesneyi) with long legs. Schwartz (1970), in dis-
cussing D. occiduus, suggested that the species crusculus-hewardi-
barbouri-occiduus might represent a phylogenetic series, despite
the interposition in this sequence of both long- and short-limbed
species. D. microblepharis stands alone; its relationships are with
the Puerto Rican D. pleei Dumeril and Bibron and the Cuban D.
delasagra Cocteau.
In the summer of 1961, while cutting bromeliads in the decid-
uous forest at the northern edge of Jamaica's Cockpit Country,
1 Miami-Dade Junior College, Miami, Florida 33167
2 BREVIORA No. 371
the extensive karst region in northwestern Jamaica, we secured a
single immature galUwasp. Despite the peculiar habitat (no Antil-
lean Diploglossus had ever been recorded from bromehads) the
Hzard bore resemblances to D. hewardi, and it was so considered
in the field. Not until 1969, when the paper dealing with D.
occiduus (Schwartz, 1970) was in preparation, was the lizard
re-examined in a routine study of D. hewardi for comparative
purposes. At that time, the difference in scutellation between D.
hewardi and the 1961 juvenile specimen quickly became appar-
ent. Further examination of the specimen indicated that, although
it resembled D. hewardi in general (being a long-limbed form),
it differed chromatically and in pattern from that species. But,
since it was immature, no further course of action was planned.
It was thus with great pleasure that I accepted the invitation
of Dr. Thomas H. Patton of the Florida State Museum to visit
Jamaica and stay at Worthy Park Estate during August 1970.
Although it was hardly likely that, even with persistent bromeliad
cutting, we would encounter another specimen of the arboreal
galliwasp, plans were made to revisit the site of capture of the
first individual (Windsor, Trelawny Parish). Thanks to the efforts
of my assistants and native help, we were successful in securing
another and adult specimen of the same form. Study of both indi-
viduals convinces me that they represent a new species, related to
D. hewardi, which has apparently taken to a bromeliad-inhabiting
niche — a niche that is virtually unoccupied by Antillean reptiles.
In the summer of 1961, I had the capable assistance of Ronald
F. Klinikowski and David C. Leber. Our activities were facilitated
by C. Bernard Lewis of the Institute of Jamaica. The 1970 trip
was made both pleasant and profitable by the presence of Dale E.
Becker, Michael T. Felix, and Danny C. Fowler, whose energy
expenditures in bromeliad cutting were noteworthy. In addition,
I have examined specimens collected by Richard Thomas in 1967,
and by Robert Brenner and Paul Moravec in the same year. All
specimens are in the Albert Schwartz Field Series (ASFS) with
the exception of the holotype of the new taxon and two speci-
mens of D. duqiiesneyi, which are in the Museum of Comparative
Zoology (MCZ) at Harvard University. All measurements are in
millimeters and color designations are from Maerz and Paul
(1950). I am especially grateful to Dr. Patton for making the
Worthy Park facilities available to us, and to Dr. Ernest E. Wil-
liams for the loan of the holotype of D. duquesneyi.
1971 BROMELIAD GALLIWASP 3
In honor of Danny C. Fowler, whose endeavors on my behalf
can only be recognized in a token fashion by associating his name
patronymically with the species, I propose that this bromeliad-
inhabiting galliwasp be called
Diploglossus fowleri, new species
Holotype. MCZ 125601, a female, from Windsor, elevation
about 500 feet (153 meters), Trelawny Parish, Jamaica, taken
15 August 1970 by Danny C. Fowler. Original number ASFS
V19902.
Paratype. ASFS 14421, same data as holotype, 12 July 1961,
D. C. Leber.
Diagnosis. An apparently moderately sized (only known adult
105 mm snout-vent length), long-limbed, bromeliad-dwelling gal-
liwasp distinguished from all other Jamaican species by a com-
bination of: 1) low number (101-103) of ventral scales between
mental and vent, 2) low number (41-43) of scales around body
at midbody, 3) angular subocular scale modally between supra-
labials 6 and 7, 4) enlarged postmental scale contacting 5 infra-
labial scales, 5) fourth toe lamellae 18-21, 6) ratio of head
width to head length high (80.0), 7) auricular opening small,
8) dorsal trunk and dorsal caudal scales keeled and striate, 9)
ventral scales smooth, 10) dorsal pattern of tans and browns
arranged in a distinct chevronate pattern, and 11) with dark
markings on the head shields.
Distribution. Known only from the type locality.
Description of holotype. An (apparently) adult female with a
snout- vent length of 105 mm and tail (almost entirely regener-
ated) 75 mm; ventral scales between mental and vent 103, 41
scales around body at midbody; fourth toe lamellae 21, angular
subocular between supralabials 6 and 7 on one side, between 7
and 8 on the other; head length 18.5, head width 14.8; ratio of
head width to head length 80.0; median enlarged postmental
(= first unpaired chin shield) small and contacting 5 infralabials.
In life, dorsal pattern consisting of a series of about 16 or 17
wood brown chevrons, their apices pointing posteriorly, from the
neck to the sacrum, on a tan ground; sides with somewhat lighter
brown continuations of these chevrons both on the neck and
between the limbs, the lateral continuations forming a series of
more or less vertical brown bars which extend ventrad to about
the level of the limb insertions; a few scattered paler tan dots or
4 BREVIORA No. 371
flecks in two vague lateral horizontal rows, associated with the
lateral brown vertical bars; head tan, with more or less symmet-
rical wood brown markings (a pair on the snout, an unpaired
median blotch in the preorbital region, a median unpaired blotch
on the posterior portion of the frontal, and the interparietal-
parietal region with the scales dark edged); a black preorbital line
on the lores; temples longitudinally streaked with very dark wood
brown; a series of three brown lines on the supralabials, one below
the eye, the two others extending vertically across the supralabials
in the loreal region, all continuous ventrally across the infralabials;
a series of three very dark brown to black nuchal blotches, the
posteriormost the largest and located above the insertion of the
forelimb; Umbs mottled brown and black dorsally, the forelimbs
additionally with some intermixed tan areas and consequently
appearing more mottled or marbled than the hindlimbs; underside
pale orange, with discrete brown longitudinal streaks (four scales
in length) or flecks on throat, and deep orange streaks on venter;
underside of limbs and tail (unregenerated portion) pale orange;
iris brown with orange pupillary ring.
Variation. The paratype is a juvenile lizard with a snout- vent
length of 66 mm. Scale counts are: 101 scales between mental
and vent, 43 scales at midbody, angular subocular between supra-
labials 6 and 7 on one side, between 8 and 9 on the other, fourth
toe lameUae 18; head length 12.7, head width 9.3 (ratio 73.2).
In hfe, the paratype was tan (PI. 14G6) dorsally with about 18
dorsal chevrons between the neck and the sacrum; the sides were
paler tan (PL 13D3). The snout was olive, with the jowls and
the base of the tail slightly reddish. The Hmbs were tan, spotted
with dark brown to black. The venter was translucent gray,
marbled with brown on the throat, and marked with reddish on
the trunk and underside of the hindlimbs and tail. The facial
markings, described for the holotype, were equally as prominent
in the juvenile paratype. The dorsal chevronate pattern was dark
brown to black, and on the sides the chevrons were continuous
with weakly defined lateral vertical brown bars, each of which
was followed by a creamy bar. Three nuchal-supra-axillary
blotches were black, and the frontal head shield had a dark ante-
rior margin with some additional dark sulTusions on the dorsal
surface of the head. The postmental scale in the paratype is very
small, but it contacts 7 infralabials; the contact on the right side
1971 BROMELIAD GALLIWASP 5
between the postmental and the third infralabiai is slight, but
the contact on the left side is slightly more broad.
Comparisons. D. fowleri needs comparison only with the three
long-limbed Jamaican species (occiduus, hewardi, duquesneyi) .
The new species differs from the short-limbed cnisculus and
barboiiri in having much larger Umbs and from microblepharis
in having the frontal longer than broad (in microblepharis, the
frontal is broader than long). Counts of ventral scales (101-103)
in fowleri overlap (of the other species) only the counts of crus-
culus (97-122, data from Grant, 1940b); all other Jamaican
species combined have ventral counts ranging from 107 to 150,
with the low count of 107 in the giant occiduus, the high count
of 150 in barbouri. In midbody scales, fowleri (41-43) overlaps
only cruscuhis (36-49) and microblepharis (43). Combined
midbody counts for all other Jamaican species (with the excep-
tion of cruscuhis and microblepharis) vary between 47 {bar-
bouri) and 59 {hewardi). In having the angular subocular
between supralabials 6 and 7, fowleri resembles barbouri and
cruscuhis but differs from hewardi and duquesneyi (7 and 8),
microblepharis (5 and 6), and occiduus (8 and 9).
In fowleri, the dorsal scales are striate and keeled; this condition
occurs in all other Jamaican galliwasps with the exception of
occiduus (dorsals striate but not keeled). In having smooth
ventrals, fowleri resembles barbouri, hewardi, occiduus, and
microblepharis. The ventral scales are striate in crusciilus and
duquesneyi. Note, however, that hewardi may have weakly striate
ventrals, and duquesneyi may have smooth ventrals. Finally, the
striate and keeled dorsal caudal scales of fowleri are Hke those of
cruscuhis and duquesneyi; all other Jamaican species have smooth
{hewardi, occiduus) or keeled {microblepharis) superior caudals.
From the above summary, it is obvious that fowleri combines
features of scuteUation of several Jamaican species in new and
different ways, and that the new species differs in combination of
these characteristics from all other Jamaican species.
Presumably, as will be noted below, D. fowleri is a local deriva-
tive of the widespread D. hewardi. The fact that the juvenile
fowleri was, in the field, considered as hewardi suggests the simi-
larities between the two species. However, in addition to the
structural differences noted in the above paragraph, the two spe-
cies differ strikingly in coloration in life and perhaps less so in
6 BREVIORA No. 371
dorsal pattern. More importantly, the size of the auricular open-
ing in fowleri is much the smaller; comparison of the opening in
the fowleri holotype and a similarly sized hewardi (ASFS 14892;
female with snout- vent length of 109 mm) reveals, even upon
casual inspection, that the auricular opening of fowleri is slightly
more than half the size of that of the hewardi. In addition, the
two species differ in that fowleri has larger ventral scales (101-
103 between mental and vent in fowleri, 113-135 in hewardi),
and fewer scales at midbody (41-43 versus 49-59 in hewardi).
The enlarged postmental contacts 7 infralabials in all hewardi ex-
amined, whereas at least in the fowleri holotype this scale con-
tacts only 5 infralabials (weak contact with 7 infralabials in the
paratype). In fowleri, the angular subocular modally hes between
supralabials 7 and 8, whereas in hewardi it lies between suprala-
bials 6 and 7. D. fowleri exceeds D. hewardi in number of fourth
toe lamellae (15-19 in 22 hewardi, 18 and 21 in two fowleri).
Finally, the head width/head length ratio in adult female hewardi
varies between 70.7 and 74.1, whereas in the fowleri holotype,
this ratio is 80.0. There are no comparably sized juvenile hewardi
for comparison of this ratio in the paratype of fowleri, but the
ratio (73.2) in this specimen lies near the upper extreme of ratios
in hewardi with shorter snout-vent lengths (45-52 mm; ratios
66.4 to 73.8).
The dorsal coloration of hewardi has been repeatedly recorded
as greenish brown to greenish tan, but some individuals have the
dorsum very dark brown (almost black) to metallic tan. The
head regularly is unmarked dorsally, and vertical subocular and
loreal lines are absent. The dorsal pattern consists of a trans-
verse series of confused bars or bar fragments; these pattern ele-
ments are usually so broken that no meaningful count can be
taken. There is a strong tendency for the hewardi dorsal pattern
elements to consist of bars, rather than chevrons as in fowleri.
Perhaps the most distinctive pattern feature of hewardi, in con-
trast to fowleri, is that of the throat. In hewardi, the throat has
a broad dark reticulum, the pattern extending as far posterior
as the forelimb insertions. This pattern is expressed even in the
smallest juveniles and becomes more intense with increasing size.
No hewardi has the discrete brown throat lines and flecks of
fowleri. In addition, the ground color of the throat in hewardi
is often blue to purplish, not pale orange as in fowleri. The deep
orange belly markings of fowleri are absent in hewardi; some
1971 BROMELIAD GALLIWASP 7
hewardi have belly markings that are not discrete as in fowled
and are gray in life.
Comparisons in detail with D. occiduus are hardly necessary.
In addition to the scutellogical differences noted above, the huge
size of occiduus (to 305 mm snout-vent length) and its presumed
terrestrial habits, coupled with its general bulk, immediately dis-
tinguish it from fowled.
The only other long-limbed Jamaican galliwasp is D. duques-
neyi. Scutellogical differences have already been noted between
this species and fowled. I have examined the holotype (MCZ
45194) and one other specimen (MCZ 45181) of duquesneyi.
The species was casually defined by Grant (1940a: 6) on the
basis of one juvenile specimen, and Cousens (1956) gave addi-
tional pigmental and pattern differences between two specimens
of duquesneyi, and hewardi. I have examined the two extant
duquesneyi and both are damaged about the body so that accurate
scale counts are difficult. Ventral scales between the mental and
vent are about 116 and 122, midbody scales are about 48 and
49, and fourth toe lamellae are 19 and 23. The angular subocular
lies between supralabials 7 and 8 on both sides of both specimens.
D. hewardi and D. duquesneyi are comparable in dorsal pattern:
the transverse markings in both are distinctly straighter and more
barlike than the chevronate pattern in fowleri. In addition, com-
parison of equally sized hewardi and duquesneyi shows that du-
quesneyi (like fowleri) has a larger auricular opening than
hewardi.
D. fowleri differs from D. duquesneyi in that the former has
fewer ventrals between mental and vent (101-103 versus 116-
122), fewer midbody scales (41-43 versus 48-49), the angular
subocular between supralabials 6 and 7 rather than between 7
and 8, and smooth rather than striate ventrals (although the holo-
type of duquesneyi has smooth ventrals). Two pattern elements
differentiate duquesneyi from fowleri: the former has the tail (at
least in juveniles) banded alternately black and sky blue (Grant,
1940b: 106), a feature unknown in any other Antillean galliwasp,
and duquesneyi has an immaculate throat and venter (color un-
known), a feature that separates duquesneyi from both fowleri
and hewardi. D. duquesneyi also lacks the prominent facial mark-
ings of D. fowleri. Finally, the head width/head length ratio (X
100) in duquesneyi is much less than this ratio in both fowleri
8 BREVIORA No. 371
and hewardi. In an apparently subadult female duquesneyi with
a snout-vent length of 96, the HW/HL ratio is 69.8, below that
of similarly sized female hewardi (ratios 70.7-13.1) and much
below that of the slightly larger female holotype of jowleri (80.0).
The HW/HL ratio in the juvenile holotype of duquesneyi (snout-
vent length about 65) is 66.4, whereas this ratio in the jowleri
paratype (snout-vent length 66) is 73.2; the HW/HL ratio in
the duquesneyi holotype falls at the lower extreme of this ratio
in smaller hewardi (ratio 66.2 to 67.3 in hewardi juveniles with
snout-vent lengths of 49 to 52).
Interestingly, Richard Thomas noted that a D. hewardi from
Darliston, Westmoreland Parish, in western Jamaica, (and far
removed from the known range of D. duquesneyi, which has been
taken only on Portland Point, Clarendon Parish, in south-central
Jamaica) with a snout-vent length of 48 mm, had pale blue distal
tail bands alternating with brown bands. There seems little doubt
that hewardi, duquesneyi, and jowleri are closely related, and
that duquesneyi and presumably jowleri are peripheral geographic
satellite species derived from parent hewardi in special situations.
Remarks. The two specimens of D. jowleri were collected
under the following circumstances. Both specimens were taken
from bromeliads along the edge of the steep trail from Windsor
Great House to Windsor Cave and thence up the escarpment of
the Cockpit Country. In the case of the holotype, a Jamaican had
been hired to cut arboreal bromeliads and had climbed a tall
tree of moderate girth (0.5 meters) just below the path in decidu-
ous forest. He had cut all but the last one or two bromeliads when
the galliwasp rapidly descended the trunk of the tree and paused
in confusion on a leaf about six feet above the ground. There is
no doubt that the animal had been disturbed from its diurnal
retreat by the chopping of adjacent bromeliads and had decided
to abandon its place of retirement. The small paratype was taken
from the moist center of a bromehad that had been growing 2.5
meters above the ground. When the bromeliad was cut and thrown
onto the narrow path, the lizard was found inside the whorls of
leaves. In both cases, the adjacent area was well forested. The
elevation along the path is about 500 feet (153 meters).
The Jamaican Cockpit Country is a karst region in north-
western Jamaica. Its extent is about 20 miles (32 kilometers)
east-west and about 10 miles (16 kilometers) north-south; the
region centers in Trelawny Parish but extends for short distances
1971 BROMELIAD GALLIWASP 9
into St. James Parish on the west and St. Elizabeth and Man-
chester parishes to the south. No roads penetrate it, but a series
of peripheral roads allows some ingress into the region. An excep-
tion is a relatively newly constructed road north of Quick Step
on the southern border of the Cockpit, where penetration of about
five miles (8 kilometers) is possible into virtually virgin territory.
In search of Sphaerodactylus and hylid and leptodactylid frogs,
we cut both terrestrial and arboreal bromeliads in several regions
associated with the Cockpit periphery and elsewhere: between
Spring Vale and Mulgrave (St. James and St. Elizabeth parishes),
between Stonehenge and Burnt Hill (Trelawny Parish); south
of Moneague on Mt. Diablo and west of Lluidas Vale (St. Cath-
erine Parish), in the Dolphin Head region between Askenish and
Town Head (Hanover and Westmoreland parishes), between
Plum Park and Garlands (St. James Parish), and between Raheen
and north of Quick Step (Trelawny Parish). In no case did we
secure D. fowleri, although a single D. cnisculus was secured from
a terrestrial bromehad north of Cave in Westmoreland Parish.
The possibiUty remains that D. fowleri is not an obligate inhab-
itant of bromeliads, and that it is a terrestrial galliwasp that, in
the pitted, pocked, and rock-strewn Cockpit Country, finds diur-
nal sanctuary in terrestrial situations from which it would be a
lucky collector indeed who would secure it. On the other hand,
there is no evidence to controvert the apparent fact that D. fowled
is indeed a bromeliad dweller and that it occurs in no other situ-
ation. If such is the case, it must be either extremely uncommon,
remarkably elusive, or ecologically or altitudinally restricted in
some presently unknown fashion. Along these lines, see Under-
wood's (1959: 1) comments on his inability to secure a second
specimen of D. microblepharis. The fact that the area where the
microblepJiaris was secured backs upon limestone hills suggests,
as Underwood stated, that it may have wandered from its usual
habitat into a situation where it was fortuitously secured with
relative ease.
Diploglossiis fowleri is not known to be sympatric with any
other species of galhwasp. However, D. hewardi has been taken
1.5 miles NW of Windsor, and D. cruscidiis has been secured 3.0
miles N W of Windsor — both in terrestrial situations. In addi-
tion, D. barbouri has been collected along the eastern margin of
the Cockpit Country between Stonehenge and Burnt Hill. The
10 BREVIORA No. 371
lack of precisely sympatric records between fowleri and any of
these three species is not surprising, since, as pointed out above,
collecting galliwasps within the Cockpit Country itself is a difficult
and well-nigh impossible task except in especially favorable local-
ities. If fowleri is truly bromeliadophilous, then it may in places
be syntopic with D. crusculus, but such syntopy remains to be
encountered.
As presently understood, then, D. fowleri is a bromehad-
inhabiting galliwasp that is presumably limited to the Cockpit
Country area and possibly to lower elevations in that region. It
is rather surprising that the bromeliad niche has been so neglected
by Antillean reptiles, in contrast to Antillean amphibians. Cer-
tainly Jamaica has the highest share of bromeUadicoles, both
amphibians and reptiles; in addition to D. fowleri, Hyla brunnea,
H. wilderi, H. marianae, and Eleutherodactylus jamaicensis are
obligate bromeliad dwellers, and several other frogs {E. grabhami,
E. cundalli, E. pantoni) are encountered with regularity in terres-
trial bromeliads. Among reptiles, Sphaerodactylus oxyrhinus
appears to be confined to this situation, and a new species of
Sphaerodactylus, to be described by Richard Thomas, likewise is
thus limited in habitat. S. argiis, D. crusculus, and Tropidophis
haetianus are encountered in bromefiads upon occasion. This list
of both obligate and facultative bromeliadicoles far exceeds that
from any other Antillean island. On the other hand, no one has
systematically cut arboreal and terrestrial bromefiads elsewhere
than on Jamaica. It seems likely that this is a niche that will
well repay investigation on other Antillean islands.
LITERATURE CITED
CousENs, Peggy N. 1956. Notes on the Jamaican and Cayman Island
lizards of the genus Celestiis. Breviora, No. 56: 1-6.
Grant, Chapman. 1940a. Notes on the reptiles and amphibians of Jamaica,
with diagnoses of new species and subspecies. Jamaica To-day. London
and Aylesbury, Hazell, Watson, and Viney, Ltd. Chapter 15: 151-157.
1940b. II. The Reptiles. In Lynn, W. G., and C. Grant,
The herpetology of Jamaica. Bull. Inst. Jamaica, Sci. Ser., 1: 1-148.
Maerz, a., and M. Rea Paul. 1950. A Dictionary of Color. New York,
McGraw-Hill Book Co., pp. i-vii, 1-23, 137-208, 56 pis.
Schwartz, Albert. 1970. A new species of large Diploglossus (Sauria:
Anguidae) from Hispaniola. Proc. Biol. Soc. Washington, 82(60):
777-788.
Underwood, Garth. 1959. A new Jamaican galliwasp (Sauria, Anguidae).
Breviora, No. 102: 1-13.
BREVIORA
Mmsemim of Contiparative Zoology
Cambridge, Mass. 31 March, 1971 Number 372
THE PALEONTOLOGY AND EVOLUTION OF CERION II:
AGE AND FAUNA OF INDIAN SHELL MIDDENS
ON CURACAO AND ARUBA
Stephen Jay Gould
Abstract. Cerion iiva has been found in great abundance in three Meso-
Indian (preceramic) shell middens on Curasao. Shells from all three sites
yield radiocarbon ages of about 4000 years B.P. Different groups of Meso-
Indians from Venezuela reached Curasao and the nearby island of Cubagua
at about the same time. A Neo-Indian (ceramic) midden on Aruba is ap-
proximately 1500 radiocarbon years old. Lists of the molluscan fauna from
all sites contain only intertidal and shallow water species. Collecting areas
can be specified by noting differences among sites in the presence of species
from various environments (rocky intertidal, mangrove, shallow grassy and
shallow rocky).
In the shell middens, Cerion presents two outstanding features: 1) almost
all shells have had the apical whorls removed artificially and, 2) shells are
larger than any living today. The apical whorls were removed by striking;
flint tools found at the sites accomplish this task easily. This was done to
release the internal vacuum and allow the animal to be sucked out through
the normal aperture. Larger shells might indicate, since modern Cerion is
so phenotypically variable, that the climate of Curasao 4000 years ago was
more moist (and therefore more hospitable) than today. But there is no
independent evidence for more rainfall at that time. If the effect is mainly
genetic, these shells might come from relict populations, adapted to the
pluvials of the previous glaciation. Cerion iiva has been found in a shell
midden in Venezuela; this establishes the reciprocity of trade between main-
land and offshore islands.
INTRODUCTION
Only a few mollusks have won entry into the Papiamento lan-
guage of the Dutch Leeward Islands. These are mostly edible
species — kiwci (Cittarium pica), karko (Strombus gigas), and
tcipa koncha ("cover shell" — a general name for chitons). Yet
2 BREVIORA No. 372
Cerion uva, the ubiquitous pulmonale of these islands, stands out
for the plethora of names attached to it, names that distinguish
small from large and beach from bush. Nevertheless, Cerion plays
almost no role in the economy of these islands today — though
one of its names, kokoUshi kalakuna (turkey shell), reflects the
fact that it is sometimes fed to turkeys as a source of lime. It
is never eaten, save as an aphrodisiac by some older residents
who believe that sea shells preserve sexual potency (and do not
realize that this halophilic pulmonale, which lives just landward
of Tectariits muricatus, does not come from the sea). But to
another people, the original Indian inhabitants of Curasao, Cerion
uva was a major source of food, for the oldest middens of the
island are crammed with their shells.
Of the many sheU sites that have been studied (Van Heekeren,
1960: 103-109, for review of archaeological work and Van Heek-
eren, 1963), Cerion is known only from the older, preceramic
middens of Curasao. Whenever it occurs, it presents two peculi-
arities: sheUs are far larger than the largest living C. uva, and
most all have had the apical whorls removed artificially.
Thanks to the kindness of Father Paul Brenneker and Mr.
Elis JuUana, local collectors, folklorists, and historians (and my
informants for the opening paragraph), and Dr. F. Creutzberg,
Director of the Biological Station at Piscadera Baai, Curasao,
I had the opportunity to study the sheU sites during the summer
of 1968. In this paper, I shall review the archaeological setting of
these islands, report on radiocarbon dating of the shell sites, tabu-
late the fauna of each and present environmental interpretations,
and discuss the occurrences of Cerion with special reference to
the peculiarities mentioned above.
CARIBBEAN PREHISTORY AND DESCRIPTION OF SITES
The Dutch Leeward Islands are tied, geographically, to Vene-
zuela. Aruba, only 27 km from the mainland, lies on the coastal
shelf, in easily navigable waters. Curacao and Bonaire are more
distant (64 and 87 km respectively), and the passage is deeper
(up to 1500 m) and more treacherous (Van Heekeren, 1960:
103). The early colonization of these islands must be discussed
in the context of Venezuelan archaeology (Cruxent and Rouse,
1958-59, 1969; Rouse and Cruxent, 1963; Rouse, 1960,
1964, 1966).
1971 CERION FROM INDIAN SHELL-HEAPS 3
The Pre-Columbian inhabitants of Venezuela and the Carib-
bean are designated Paleo-, Meso-, or Neo-Indians on the basis
of technology and inferred economy. Although the three stages
do express a chronological progression, none of their artifacts
function as "index fossils" in establishing contemporaneity
throughout the Caribbean, for the traits of a new stage are
attained at different times by different peoples. There were, for
example, still some preceramic Meso-Indians on Haiti and Western
Cuba when Columbus arrived (Rouse, 1966).
The original inhabitants of the New World were Paleo-Indians,
"hunters of mammoths and other large land mammals" (Rouse,
1966: 125). Their stone tools have been found in Venezuela and
designated as markers of the Joboid Series. They date, approxi-
mately, from 17,000-7,000 B.P. The oldest radiocarbon date
for Joboid charcoal is 16,870 years B.P. (Rouse and Cruxent,
1963). In earlier works, Cruxent and Rouse held that Paleo-
Indians were not sea-farers, but Paleo-Indian sites have recently
been found at Mordan in the Dominican Republic and dated to
at least 4560 radiocarbon years B.P. They beheve, moreover,
that the Mordan site is predated by another at Casimira that may
be as much as 7,000 years old (Cruxent and Rouse, 1969).
Although the mainland source of these first Hispaniolans is not
known, these finds indicate that some Paleo-Indians crossed con-
siderable stretches of ocean, probably on rafts and by accident
(Cruxent and Rouse, 1969).
Much scholarly agitation of late has been directed to the issue
of whether or not Paleo-Indians were responsible for the extermi-
nation of large land mammals (Martin and Wright, 1967). In
any event, their demise drew our pre-agricultural people to the
sea and inaugurated MesoTndian culture, characterized by "rela-
tively few stone tools. Projectile points are made of bone rather
than stone and shell artifacts are common, reflecting the mari-
time orientation" (Rouse, 1966: 126). Meso-Indian artifacts in
Caribbean Venezuela belong to the Manicuaroid Series and date,
approximately, from 7,000 to 3,000 years B.P. The oldest radio-
carbon date for mainland Venezuelan Meso-Indians is 5750 B.P.
(Rouse and Cruxent, 1963). There is an extensive Meso-Indian
site on Cubagua, another of Venezuela's offshore islands. Char-
coal from the base of this deposit dates at 4275 radiocarbon
years B.P.
4 BREVIORA No. 372
The subsequent Neo-Indian culture is "marked by pottery
making and fully developed agriculture" (Rouse, 1966: 126).
The invention of pottery was the crucial archaeological event that
inaugurated the Neo-Indian period; therefore MesoTndian and
earlier sites are often designated simply as "preceramic." Agri-
culture, with manioc as a staple crop, and pottery were developed
in the Orinoco Valley during the 2nd millennium B.C. During
the 1st millennium B.C., some Neo-Indians moved out to the
coast and became sea-farers. Displacing Meso-Indians as they
went, they migrated to the coastal islands, up the Lesser Antilles
and reached the Greater Antilles ca. 250 A.D. and the Bahamas
ca. 1000 A.D. This displacement was still occurring when Colum-
bus reached the New World (Cruxent and Rouse, 1969).
The Cerion sites of Curasao are all Meso-Indian in nature. I
studied the following three sites:
1. Rooi Rincon — North coast, west of Hato Airfield; in soil
at the base of a small cave in a raised Pleistocene reef that also
houses the larger cavern of Hato and several others; approximately
40 m above present sea level and 1 km from the coast. This well-
known site was excavated by Cruxent in 1965 (Tamers, 1967)
and by Van Heekeren in 1960 (Van Heekeren, 1963). Crudely
chipped stone tools and flint flakes are common but, after dig-
ging for 14 days. Van Heekeren found only one other artifact, a
shell disc bead (Van Heekeren, 1963: 5). The naturally broken
columellar tips of Strombus gigas are similar in form to some of
the fashioned shell gouges common in the Manicuaroid deposits
of Cubagua (Cruxent and Rouse, 1958-59); they may have been
used for digging meat out of sheUs. Many other natural objects
could have been used as tools. Particularly suspect are the
smoothly eroded and fairly pointed branches of the stag horn
coral, Acropora cervicornis, that are fairly common at this site
and at Kintjan (site 2). These, obviously, have no nutritional
value and must have been carried to the site for some other pur-
pose. Other objects, land crab claws for example, might have
been used for digging meat from shells after their own contents
had been consumed. I found a few bits of charcoal: some of
the shells are strongly scorched. Cruxent says of this deposit:
"A Meso-Indian complex of collectors with industry of stone
chips. Classified as a marginal development of El Jobo. No
archaeologic station of this type presently known in Venezuela"
{in Tamers. 1967: 244).
971
CERION FROM INDIAN SHELL-HEAPS
2. Kintjiin — Near south coast, east ^ Willemstad. The area,
a hillslope, is being cleared for construction and shells are loose
at the surface; their presence in a small area indicates original
concentration in a coherent deposit. Flint chips and crude stone
tools are, as at Rooi Rincon, common at this site.
3. Tafelberg — Near south coast, just east of the Tafelberg
Santa Barbara. Only a few shells could be collected from the
recently blasted rubble of these phosphate workings. Mr. Harry
Evers, engineer at the Tafelberg phosphate workings, informs me
that, prior to the blasting, the shell heap was a coherent deposit
with two layers, marine shells at the base and decapitated Cerion
at the top. I found no artifacts at this much disturbed site.
Dr. P. Wagenaar Hummelinck, pre-eminent natural historian of
these islands, has told me (personal communication, 1970) of
one additional Cerion locality at Hato Cave; I have not seen this
site. He also states that he knows of no other Cerion site on any
of the three islands.
For comparison, I add to the Cerion sites of Curacao one
later, Neo-Indian deposit from Aruba:
t:.t V-
^pt>^^
Figure 1. Artifacts from Ceru Canashito, Aruba.
la) left: rock drawing, presumably depicting a pregnant woman,
lb) right: shell disc made from Melongena melongena. Actual
height: 43 mm.
6 BREVIORA No. 372
4. Ceru Canashito — North slope of this Hmestone terrace.
I chose this among the many Neo-Indian sites of Aruba for two
of its outstanding features. Good skeletal material has been col-
lected from the caves near its summit (Tacoma, 1959), and these
caves contain some of the best of the celebrated and mysterious
rock paintings of these islands (Hummelinck, 1953, 1957). One
of these, probably depicting a pregnant woman, is reproduced as
Figure la. (There is, of course, no reason to assume that the
rock drawings are contemporaneous with the shells; Van Heek-
eren (1960), in fact, suspects that they were fashioned by Meso-
Indians and venerated by later inhabitants.) Shells occur at all
levels of the slope, but are concentrated by gravity at the base
in an inhomogeneous deposit. Sherds of a coarse, unornamented,
grit-tempered pottery are common. Shell artifacts include the
columellar points of Strombus gigas and the unperforated shell
disc, made from the outer whorl of Melongena melongena, shown
in Figure lb. Such unperforated shell discs are common on the
islands; their function is unknown (Van Heekeren, 1960: 112).
AGE OF THE SHELL MIDDENS
Tamers (1967) reported the first radiocarbon dates from
archaeological sites in the Dutch Leeward Islands; all samples
were charcoal and all were supplied by Cruxent. Included are
five dates for the Rooi Rincon shell midden, two from a pit previ-
ously excavated by Van Heekeren and three from two new pits.
The dates range from 3900 ± 50 to 4490 ± 60 with a mean of
4194 radiocarbon years (see Stuiver and Suess, 1966 on the
relationship between radiocarbon and calendar years). These
are the only dates previously calculated for preceramic sites on
these islands.
Radiocarbon ages were determined for 1 1 shell samples by
Geochron Laboratories, Inc., Cambridge, Massachusetts (Chama
macerophylla and Cittarium pica from each of the five sites and
Anadara notob'iUs from Ceru Canashito) . "The shells were cleaned
of foreign material and were thoroughly leached with dilute HCl
in an ultrasonic cleaner to remove the surficial layer of carbonate
and expose fresh material. The cleaned shells were then hydro-
lyzed to recover CO., for the analysis" (personal communication
from H. W. Krueger of Geochron). Dates are based on a half-
life of 5570 years and referenced to 1950 A.D.
1971 CERION FROM INDIAN SHELL-HEAPS 7
Dates based on shells are not as reliable as those determined
for pure carbon (charcoal), for CaCO,, is often altered by per-
colating, acidic groundwaters. I was anxious to determine the
correspondence between shell and charcoal dates for Rooi Rincon;
I found no charcoal at any of the other sites. All dates are shown
in Table 1.
The correspondence at Rooi Rincon is satisfactory, and all pre-
ceramic sites of Curacao are about 4000 radiocarbon years old.
This date is particularly interesting since it corresponds so well
with the base of the great Meso-Indian site at Punta Gorda,
Cubagua Island (p. 21). The artifacts of this Cubagua complex
of the Manicuaroid series differ greatly from those of Rooi Rin-
con (Cruxent and Rouse, 1958-59) and we must assume that
different groups of Meso-Indians from Venezuela colonized the
coastal islands at about the same time.
The great spread of dates for the Neo-Indian site of Ceru Cana-
shito can be explained in two ways. It is a very inhomogeneous
deposit of shells artificially concentrated at the base of a slope
and may represent a long span of habitation. Alternately, the Cit-
tarium date could be spuriously young. Cittarium has been and
remains a staple food of the islands. The kiwa is sold at all native
market places; shells are carried and discarded all over the island.
If this date has been falsified by the inclusion of a fairly modem
shell, then the Canashito midden may represent a more coherent
deposit, about 1500 radiocarbon years old.
FAUNA OF THE SHELL MIDDENS
In presenting these faunal lists, I have excluded the micro-
molluscs that could have played no role in the economy of the
Indians (though Tnmcatella and other rissoids are reasonably
common as accidental transports). In each site, there are a few
species that clearly dominate; these are merely listed as common.
Numbers of specimens are given for other species. I have used
Warmke and Abbott (1961) and Coomans (1958) as guides to
identification; order of listing and family allocations follow the
former source.
1. Rooi Rincon
AMPHINEURA
A cant ho pleura graniilata — common
8 BREVIORA No. 372
GASTROPODA PROSOBRANCHIA
Trochidae
Cittarium pica — common
TURBINIDAE
Astraea tecta — 1
Astraea tuber — 1
Neritidae
Merita peloronta — 11
Nerita versicolor — 6
Nerita tesselata — 4
LiTTORINIDAE
N odilittorina tuberculata — 4
Echinus nodulosus — 1
Tectarius muricatus — 1 0
Vermetidae
Petaloconchus mcgintyi — 3
Strombidae
Strombus gigas — 4 apices and 3 columellas
MURICIDAE
Murex brevifrons — 8
Magilidae
Coralliophila abbreviata — 2
Coralliophila caribbea — I
Fasciolariidae
Leucozonia nassa — 1
Xancidae
Vasum capitellum — 1
GASTROPODA PULMONATA
Cerionidae
Cerion uva — common; 18 of 129 specimens have in-
tact apices
BIVALVIA
Arcidae
Area zebra — 12 valves
Area imbricata — 4
Anadara notabilis — 4
Mytilidae
Brachidontes exustus — 2
Pteriidae
Pinctada radiata — 1 3
1971 cerion from indian shell-heaps 9
Pectinidae
Pecten ziczac — 2
LiMIDAE
Lima scabra — 10
OSTREIDAE
Ostrea jrons — 1 2
Crassostrea rhizophorae — 9 .
Chamidae
Chama macerophylla — common
Pseudochama radians — 2
Nonmolluscan remains: a few branches of stag-horn coral
{Acropora cervicornis) , land crab claws (common), a few barna-
cles, fish bones and a small fragment of an echinoderm test.
Not all these animals were eaten. Many, especially among the
snails, are small and rare at the site (turbinids, magilids, fasci-
olariids, and xancids); others (Petaloconchus and barnacles)
cement to other shells and surely won a free ride on their edible
hosts (probably Chama).
The main food sources were the land snail Cerion, land crabs,
intertidal chitons, the intertidal and just subtidal snail Cittarium
and the shallow water clam, Chama; all are very abundant and
easily gathered. Less common but still important as food sources
are the conch Strombus gigas, Nerita peloronta, and Murex brev-
ifrons among the snails (the last two artificially broken in char-
acteristic ways — Figs. 2 and 3 ) and arcids, oysters, and limids
among the clams.
The shells provide an excellent picture of the environment from
which they were gathered. All the major intertidal rock-clingers
are represented (all three common West Indian Nerita, chitons,
and the famous homeomorphic series Nodilittorina-Echinus-
Tectarius). These species inhabit rocky shores in areas of active
surf. All other species can be found in less than 10 feet of water
on a varied bottom containing reefy and rocky areas (Chama,
Area, Lima) and stretches of sand and grass (Anadara, Strom-
bus). There may have been a lagoon with mangroves nearby,
for many important elements of the mangrove-root community
are present (Murex brevijrons, Ostrea jrons, Crassostrea rhizo-
phorae, and Brachidontes exustus).
Van Heekeren (1963) stated, correctly no doubt, that the
shells were collected on the nearby north coast (Fig. 4b). Since
10
BREVIORA
No. 372
Figure 2. Miire.x brevifroiis shells from Kintjan (left) and Rooi Rincon
(right). Note characteristic breakage pattern in both. This can be achieved
by placing the shell face down upon its aperture and striking the apex.
Actual height of Kintjan specimen: 54 mm.
Figure 3. Neritids from Rooi Rincon broken in characteristic fashion.
Left: apertural portion from rear; Right: apertural portion from front.
Such a break is made by placing the shell face down upon its aperture and
striking the body whorl with a blunt object. This is also the natural break-
age pattern in most cases. Right-hand fragment is 18 mm high.
1971 CERION FROM INDIAN SHELL-HEAPS 11
the unremitting trade winds blow against this coast (producing
a strong surf most unconducive to shell gathering), Van Heekeren
suggested that sea level at this earlier time was 6-7 m higher
than today. This would submerge the extensive raised reef that
forms the lower terrace all around Curacao and produce a broad
area of calmer, shallow water. (And from the supposed extent
of this change in level, he postulated a great age for the deposit
and classified it, tentatively, as Paleo-Indian.) This hypothesis
of a major shift in sea level is unnecessary for two reasons: 1)
With an age of 4000 radiocarbon years, any eustatic fall in
level is ruled out; if anything, mean sea level then was a bit
lower than today (Redfield, 1967; Milhman and Emery, 1968).
This leaves tectonic uplift. Curacao has, indeed, been uplifted
during the Pleistocene (the oldest terrace, atop the Tafelberg,
lies at 140-200 m, but 7 m in 4000 years is not likely). 2) The
trade winds do produce a strong surf along the north coast. But
Rooi Rincon lies on that part of the coast that runs due east-
west; here the winds run along the coast and the waters are fairly
calm. Modern Cerion populations illustrate the climatic results
of changes in coastal direction. Cerion lives atop the first terrace
all along the coast. In areas continually buflfetted by the strong
dry wind, they aestivate for much of their lives and remain small
as adults; they grow bigger in calmer areas. A graph of Cerion
size vs. distance from Westpunt (Fig. 4a) is a good map of coastal
direction (Fig. 4b). Cerion is small where the coast runs north-
south and large where it runs east-west. They reach their greatest
size at Rooi Rincon. Thus, Rooi Rincon lies in the only area of
Curasao that provides good conditions for shell gathering on the
north coast.
2. Kintjan
GASTROPODA PROSOBRANCHIA
Trochidae
Cittarium pica — common
Strom BiDAE
Strombus gigas — common
Cymatiidae
Charonia variegata — 1
MURICIDAE
Murex brevijrons — 3 (broken as at Rooi Rincon,
Fig. 2)
12
BREVIORA
No. 372
Melongenidae
Melongena melongena — 2
GASTROPODA PULMONATA
Cerionidae
Cerion uva — common, 7 of 347 specimens have in-
tact apices
BIVALVIA
Arcidae
Area imbricata — 13
Barbatia cancellaria — 3
Anadara notabilis -
Pteriidae
Pinctada radiata — -
Pectinidae
Pecten ziczac — 7
Limidae
Lima scabra — 6
common
Figure 4. Correlation of coastal direction and shell size.
4a) left: map of Cura^,ao. 1. Rooi Rincon at point where coast
runs east-west. 2. Kintjan. 3. Tafeiberg. 4. Schottegat (where shells at
Kintjan were collected).
1971
CERION FROM INDIAN SHELL-HEAPS
13
OSTREIDAE
Ostrea frons — 7
Crassostrea rhizophorae — 6
Chamidae
Chama macerophylla — common
NonmoUuscan remains: branches of stag-horn coral (Acropora
cervicornis), barnacles, and fish bones.
The shallow water fauna of Kintjan is very similar to that of
Rooi Rincon, both in species composition and order of dominance
{Chama and Cittarium followed by S trombus, arcids, oysters, and
limids). Since shells are not so common at Kintjan, several spe-
cies, rare and unimportant at Rooi Rincon, are not found here.
10 20 30
Distance from Westpunt (miles)
4b) right: Mean shell heights (20 adults per sample) for local
populations living in similar microhabitats directly on the first terrace along
the east coast of Curasao. Shells are largest where trade winds do not hit
coast directly.
14 BREVIORA No. 372
I found no land crabs at Kintjan, but Cerion uva is even more
common here than at Rooi Rincon. There is, however, one out-
standing difference between the two sites: there are no intertidal
rock-dwellers at Kintjan (neritids, littorinids, or chitons), while
all the common forms are found at Rooi Rincon. This difference
permits us to specify the collecting area for Kintjan shells.
The entire periphery of Curasao is framed by an uplifted Pleis-
tocene reef; intertidal forms are common all around the coast.
But the central areas are underlain by volcanic rocks that erode
more easily than the coastal limestone. During the last glacial
period, when sea levels were lower, extensive drainage systems
were developed on the volcanic terrain; these breached the harder
limestone rim in only a few places. These valley systems were
drowned when sea level rose and produced the outstanding pro-
tected harbors that characterize all three islands: narrow inlets
with expansive inland waters. Willemstad, the capital of Curacao,
is built on both sides of the largest harbor, the Schottegat. The
inland shores of the Schottegat are volcanic; in the absence of
strong surf and a rocky coast, the rock-dwelling intertidal forms
do not inhabit these shores. I conclude that the Kintjan shells
were collected in the Schottegat (Fig. 4b); the extensive, calm,
shallow waters provided an excellent site for gathering.
3. Tafelberg
GASTROPODA PROSOBRANCHIA
Trochidae
Cittarinm pica — several fragments
LiTTORINIDAE
Tectarius muricatus
GASTROPODA PULMONATA
Cerionidae
Cerion uva — common, 5 of 1 1 1 have intact apices
BIVALVIA
Arcidae
Area imbricata — 1
Chamidae
Chama maeerophylla — common
The site has been thoroughly disturbed by blasting.
1971 CERION FROM INDIAN SHELL-HEAPS 15
4. Ceru Canashito
AMPHINEURA
Acanthopleiira granulata — 4 plates
GASTROPODA
Trochidae
Cittarium pica — 8
Turbinidae
Astraea tecta — 1
Neritidae
Nerita tessellata — 3
Littorinidae
Tectarius miiricatiis — 5
MODULIDAE
Modulus modulus — 1
Cerithiidae
Cerithium algicola — 1
Cerithium Utteratum — 1
Strombidae
Strombus gigas — common
Muricidae
Murex pomum — 7
Murex brevijrons — 1
Thais deltoidea — 1
Melongenidae
Melongena melongena — common
Xancidae
Vasum muricatum — common
BIVALVIA
Arcidae
A nadara notabilis — common
Pteriidae
Pinctada radiata — 1
LUCINIDAE
Codakia orbicularis — common
Chamidae
Cliama macerophylla — common
Pseudochama radians — 1
16 BREVIORA No. 372
Intertidal rock-dwellers are found here, but the series is not
nearly so complete as at Rooi Rincon (only one Nerita, Tectarius,
but neither Echinus nor Nodilittorina). Among shallow water
forms, there are two major differences between Canashito and
both Rooi Rincon and Kintjan. The Curasao sites contained a
suite of mangrove-dwellers that are completely absent here (Cana-
shito yielded one Murex brevijrons, a common mangrove form,
but Murex pomum, an open water species absent from both Cura-
sao sites, is the common Murex here). In addition, Canashito
contains a suite of shells {Modulus, the two Cerithium species
and, especially, the common Codakia orbicularis) that inhabit
grass and algal beds; none of these occur in the Curasao deposits.
The shells were probably collected in calm waters off the leeward
south coast, near the site of the present airport.
CERION UVA IN THE PRECERAMIC MIDDENS
OF CURACAO
In all three preceramic middens of Curagao, the most common
moUuscan shell is that of the land snail Cerion uva. These shells
present two outstanding features: more than 80 percent in each
locality have lost their apical whorls and shells are larger and
more variable than modern specimens.
1. Removal of the apical whorls. By reason and experiment,
one of a list of possible proposals can be identified as the cause
of removal. I list the suggestions made to me by many friends
and colleagues.
A) Natural removal
B) Artificial removal
i) by biting
ii) by rubbing
iii) by crushing (striking with the shell held up-
right)
iv) by slicing (striking with the shell placed on
its side).
Although the apical whorls form the weakest part of the shell,
I do not believe that they could have been lost naturally by so
many specimens. I have extensive collections of much older fos-
sils from fissure-fills on Aruba. These tumbled, often down sev-
eral meters, into the fissures, suffered strong compaction, under-
went tectonic uplift and still retain, in almost all cases, the apical
971
CERION FROM INDIAN SHELL-HEAPS
17
whorls. I have never seen a natural accumulation, cither recent
or fossil, in which many specimens are missing their apical whorls.
After suffering one dental misfortune, I am quite sure that the
tops cannot be bitten off. Apices can be removed by rubbing
either against limestone or volcanic rock, but the process is
much too laborious and time-consuming. I am convinced that
the tops were removed by striking. They were not crushed by
striking the top of the shell while holding the bottom against a
substrate (and keeping the shell vertical), for this process invari-
ably breaks the lower lip of the aperture before crushing the top.
If, however, the shell is placed on its side, horizontally against
the substrate, the top can easily be removed by striking with a
sharp instrument. In fact, the flint chips and stone tools of Rooi
Rincon and Kintjan, are excellent devices for this purpose. With
a bit of practice, the apices can be removed with a single blow.
This leaves open the question of why the apices were removed.
1 can imagine three interpretations:
A) Removal is unrelated to eating; the shells were used for
an ornamental or other purpose.
B) When the top is removed, the animal can be sucked out
through the apical hole thus produced.
C) Removal of the top aids, somehow, in sucking the animal
out through its normal aperture.
I cannot imagine what nongastronomical purpose so many
thousand decapitated shells could have served. Moreover, the
following demonstration that decapitation is an aid to removal
of the animal argues strongly against A.
Figure 5. X-ray photographs of decapitated Ccrion uva from Kintjan
(left 2 specimens) and Rooi Rincon (right 2). Since internal whorl parti-
tions are intact, animal was not removed through apical hole. Specimen on
left is 32.8 mm high.
18 BREVIORA No. 372
If the animal were sucked out through the top, some of the
internal whorl partitions would have to be broken, for the large
foot could not fit in the small whorls left near the top of the shell.
X-ray photographs of decapitated shells (Fig. 5) show clearly
that the whorl partitions are never disturbed. The animal could
not have been extracted through the apical hole.
If you take an intact shell with its animal inside and suck as
hard as possible at the aperture, the animal cannot be extracted.
But, when the apex is removed, a single hard suck upon the aper-
ture will extract either the large foot of the animal or the entire
body itself. Removal of the top breaks the vacuum inside the shell
and facilitates the extraction of its contents. The entire process
is really quite efficient: one strike, one suck, and the animal is
removed. Several can be eaten in a minute (though I recommend
Cerion only to the starving).
Somehow, I find it satisfying to think that the Meso-Indians of
Curacao discovered an important physical principle for such a
practical procedure. This idea, so obvious to all of us who were
raised in the pre pop-top age of the beer can industry, is by no
means a self-evident principle.
2. Variation and jorm of Cerion uva. Any sample from a shell
midden is, of course, strongly biased from a biometrical point of
view. The probable bias, in these cases, is twofold: the selection
of large individuals (for Cerion is not a large snail and much
work must be expended for little nutrition), and the amalgama-
tion of shells from several local populations.
Much has been made in the literature of the extreme intraspe-
cific variability of land snail shells. This indeed is true, but it is
usually of a particular kind (and this is rarely emphasized). The
variation is interpopulational, i.e., the shells of any local popu-
lation are not unusually variable, but differences among the means
of local populations are often extreme. Thus, it is likely that our
two biases will afi'ect the mean of a midden sample in opposite
ways: the selection of large shells will augment the mean, but
the amalgamation of large individuals from several local popula-
tions will produce a midden mean smaller than the true mean of
a local population with large shells.
The rise in variability from amalgamation of local populations
can be gauged by comparing coefficients of variation (C.V.)
(Simpson, Roe, and Lewontin, 1960: 89-95) of midden samples
1971
CERION FROM INDIAN SHELL-HEAPS
19
and modern local populations for the same character. Table 2
presents C.V/s for shell height of the three midden samples and
a mean value for 69 modern local populations (Gould, unpub-
lished data for monograph in preparation; N = 20 for all samples,
midden and modern; values for midden shells are estimates for
actual height with decapitated apical whorls restored; all shells
are adults with completed growth ) . All midden means are above
the modern grand mean. Rooi Rincon and Tafelberg are within
the span of modern C.V.'s (4.03 to 10.18), but, at 15.45, shells
from Kintjan are far more variable than those of any modern
local population.
The striking feature of midden samples is the large size of some
of their shells. Fortunately, Cerion iiva is among the world's best
known land snails from a biometrical point of view. Three major
studies have been done in this century: by Baker in the earlv
1920's (Baker, 1924), by Hummelinck in the late 1930's (Hum'-
mehnck, 1940) and by myself during the past two years. Table
2 compares the heights of shells in midden and modern samples.
Each modern study has uncovered a local population with greater
mean height than the smallest midden sample, and one of Hum-
melinck's local populations exceeds the largest midden sample in
mean height. Still, of course, the midden means are all well above
T^'*'
Figure 6. Comparison of largest shell heap (left, from Kintjan, 34.3 mm
high) Cerion and largest modern shell. Difference is much more striking in
actual shells in which areal artifact of two dimensional representation is
lost and judgment of size is made more properly by volume.
20 BREVIORA No. 372
the grand mean of means for each modern study. However, as
mentioned previously, the midden means are almost surely lower
than the true means of local populations with large shells living at
that time. A more appropriate comparison might be made using
maximal size.
Among almost 12,000 modern snails from 248 local popula-
tions over 50 years, no snail greater than 30 mm in height has
ever been found. (In only one of Hummelinck's local populations
did any individuals exceed 29 mm; neither Baker nor Gould found
any taller than 28.5 mm.) Yet snails exceeding 30 mm in height
are very common in two of the three midden samples and, at 34.3
mm, the largest snail from Kintjan dwarfs my modern "giant"
(Fig. 6).
Two separate factors can make a snail tall, and both operated
to produce the large midden shells. First, a snail can increase in
height simply by adding more whorls. Each of the decapitated
shells of Figure 5 shows 1 1 whorls below the break; the complete
shell would have had one or two more postprotoconch whorls.
Modern shells with more than IOV2 postprotoconch whorls are a
great rarity (Baker and Hummelinck included protoconch whorls
in their count, hence their larger figures). Secondly, a tall snail
may have as many whorls as a smaller one, but simply have larger
whorls. Protoconch size is a good measure of general whorl size
(Gould, 1969). Only Rooi Rincon has enough complete shells
to permit the calculation of mean protoconch width. At 1.67 mm,
mean protoconch width for Rooi Rincon is at the top of the range
of modem mean widths (1.41-1.69 mm for 69 samples, N ^ 20
for each sample ) . The midden shells grew more whorls than any
modern sample and had larger whorls than most.
Why were the midden snails larger than modern snails? All
three modern studies have demonstrated the extreme phenotypic
plasticity of Cerion iiva. Shell size of adults is a direct function of
microenvironment; snails are large when habitats are moist, calm,
and well vegetated. Curasao today is an arid island. It receives
only 17-22 inches of rain per year, most in brief downpours. It
is hard to imagine a less hospitable area in the West Indies for
pre-agricultural Meso-lndians. I do not know what they could
have found, in this cactus-covered land, to supplement a diet of
sea food. It is therefore tempting to think that the large midden
shells indicate a wetter climate that might have supplied to Meso-
lndians some of the tropical fruits that adorn most West Indian
1971 CERION FROM INDIAN SHELL-HEAPS 21
islands. Unfortunately, there is no other evidence for greater rain-
fall 4000 years ago. If Curasao were much larger or higher than
it is today, continental effects might lead to increased rainfall.
But the eustatic rise of sea level has not been more than 10 feet
during the past 4000 years (Redfield, 1967; Milliman and Emery,
1968) and the direction of tectonic movement has been upward
(Weyl, 1966). Rouse and Cruxent (1936: 38) believe that tem-
peratures and rainfall have not varied appreciably during the past
5000 years in Venezuela and surrounding areas.
If large size is not an immediate phenotypic response to local
conditions more favorable than today's, then I suspect that the
midden snails were programmed to be large, i.e., that the effect
is mainly genetic. In this case they probably represent the relict
populations of snails that had been genetically adapted to more
favorable conditions during pluvial cycles of the previous glacial
period. In any event, they served the Meso-Indians well; it would
be hard to make a meal of modern Cerion.
There is an interesting postscript to the relationship of Cerion
with Meso-Indians. There is considerable evidence for trade
between the mainland and coastal islands, but it is all unidirec-
tional. Rouse and Cruxent (1963: 45) found trade pottery from
Venezuela in the Punta Gorda complex of the Manicuaroid Series
on Cubagua. Du Ry (1960: 85) discovered that the oldest pot-
tery of Aruba is finer in texture than later examples. He assumes
that this first pottery was imported from northeastern Venezuela
and that the later work is indigenous. In a nearly-forgotten work,
Berry (1934) found Cerion iiva in an Indian shell heap near Lake
Valencia, Venezuela. Berry was not convinced that these shells
were imported from the Dutch Leeward Islands. But his argu-
ment that Cerion might have once inhabited the shores of Lake
Valencia can be discounted because this halophile would not sur-
vive so far inland. I also doubt that Cerion inhabited the coast of
Venezuela, for it has never been recorded from shell heaps there.
Since there is no evidence that Cerion uva ever lived elsewhere
than the Dutch West Indies, I conclude that the Valencia speci-
mens establish the reciprocity of transport between Venezuela
and the islands.
22 BREVIORA No. 372
ACKNOWLEDGEMENTS
I thank Father Paul Brenneker and Mr. Ehs Juhana, local folk-
lorists and archaeologists, who collected with me at Rooi Rincon;
Dr. F. Creutzberg, Director of CARMABI, who showed me the
Kintjan site, and Mr. Harry Evers, who allowed me to collect at
the Tafelberg midden. This work was supported, in part, by
N.S.F. Grant No. GB-12553.
1971 CERION FROM INDIAN SHELL-HEAPS 23
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LITERATURE CITED
Baker, H. B. 1924. Land and freshwater molluscs of the Dutch Leeward
Islands. Occas. Pap. Mus. Zool. Univ. Mich., No. 152. 158 pp.
Berry, C. T. 1934. Pleistocene remains found near Lake Tacarigua, Vene-
zuela. J. Washington Acad. Sci., 24: 387-395.
CoOMANS, H. E. 1958. A survey of the littoral Gastropoda of the Nether-
lands Antilles and other Caribbean Islands. Stud. Fauna Curasao,
8: 42-111.
Cruxent, J. M., AND L Rouse. 1958-59. An Archaeological Chronology of
Venezuela. Social Sci. Monogr. VI, vols. I, 277 pp. and 11, 223 pp.
Pan American Union, Washington, D.C.
1969. Early man in the West Indies. Sci. Am. 421: 42-52.
Du Ry, C. J. 1960. Notes on the pottery of Aruba, Curasao, and Bonaire.
Stud. Archaeol. Netherlands Antilles, I: 81-102.
Gould, S. J. 1969. An evolutionary microcosm: Pleistocene and Recent
history of the land snail P. (Poecilozonites) in Bermuda. Bull. Mus.
Comp. Zool., 138: 407-532.
Heekeren, H. R., VAN. 1960. A survey of the non-ceramic artifacts of
Aruba, Curacao, and Bonaire. Stud. Archaeol. Netherland Antilles, II:
103-120.
1963. Prehistorical research on the islands of Curagao, Aruba,
and Bonaire m I960. Stud. Archaeol. Netherlands Antilles, III: 1-24.
Hummelinck, p. W. 1940. Mollusks of the genera Cerion and Tiidoni.
Stud. Fauna Curasao, Aruba, Bonaire, and the Venezuelan Islands, 2:
43-82.
1953. Rotstekeningen van Curasao, Aruba en Bonaire.
(Linear rock designs of Curasao. Aruba, and Bonaire). West-Indische
Gids, 34: 173-209.
1957. Rotstekeningen van Cursn^diO, Aruba en Bonaire. West-
Indische Gids, 37: 93-126.
Martin, P. S., and H. E. Wright, Jr. (eds.). 1967. Pleistocene Extinctions:
the Search for a Cause. New Haven, Yale Univ. Press. 453 pp.
Milliman, J. D., and K. O. Emery. 1968. Sea levels during the past 35,000
years. Science, 162: 1121-1123.
Redfield. a. C. 1967. Postglacial change in sea level in the western North
Atlantic Ocean. Science, 157: 687-691.
Rouse, I. 1960. The entry of man into the West Indies. Yale Univ. Publ.
Anthrop., No. 61. 26 pp.
1964. Prehistory of the West Indies. Science, 144: 499-513.
1966. Paleo- and Meso-Indians of the Caribbean area. Qua-
ternaria, 8: 125-132.
26 BREVIORA No. 372
Rouse, I., and J. M. Cruxent. 1963. Venezuelan Archaeology. New
Haven, Yale Univ. Press. 179 pp.
Simpson, G. G., A. Roe, and R. C. Lewontin. 1960. Quantitative Zoology.
New York, Harcourt, Brace and Co. 440 pp.
Stuiver, M., and H. E. Suess. 1966. On the relationship between radio-
carbon dates and true sample ages. Radiocarbon 8: 534-540.
Tacoma, J. 1959. Indian skeletal remains from Aruba. Stud. Phys. Anthrop.
Netherlands Antilles, II: 95-112.
Tamers, M. A. 1967. Instituto Venezolano de Investigaciones Cientificas.
Natural radiocarbon measurements 111. Radiocarbon, 9: 237-245.
Warmke, G. L., and R. T. Abbott. 1961. Caribbean Seashells. Livingston
Co. 346 pp.
Weyl, R. 1966. Geologic der Antillen. Berlin, Gebriider Borntraeger.
410 pp.
BREVIORA
Mnasemim of Comiparative Zoology
Cambridge, Mass. 31 March, 1971 Number 373
THE CHANARES (ARGENTINA) TRIASSIC
REPTILE FAUNA.
VIII. A FRAGMENTARY SKULL OF A LARGE THECODONT,
LUPEROSUCHUS FR ACTUS
Alfred Sherwood Romer
Abstract. Incomplete remains of a large skull, not improbably repre-
S2nting a raiiisuchid thecodont from the early Middle Triassic of Argentina,
are described as Luperosiichiis fractns, gen. et sp. nov. Large dermal scutes,
found isolated, may pertain to this form.
INTRODUCTION
A moderate number of specimens assignable to the reptilian
order Thecodontia are present in our Chaiiares collections. Apart
from materials that are difficult of interpretation or assignment,
there are definitely present: ( 1 ) a small and primitive member
of the Ornithosuchidae; (2, 3) two long-snouted forms, with gen-
eral proportions resembling crocodilians, but without any positive
indications of affinities with that group; (4) a small form with a
very lightly built skull, of which the limbs are unknown; (5, 6)
two forms known from very slender hind limbs of unusual con-
struction; and (7) a large animal, probably a rauisuchid. repre-
sented only by a partial skull. In addition there are various
isolated materials, pseudosuchian in nature. In the present short
paper 1 shall describe only the last specimen listed, leaving the
others for later description.
Since our collections were made, Sr. Bonaparte of Tucuman
has made several visits to the Chaiiares region, and found, inter
alia, a fair amount of thecodont material. He invited me to make
use of this in my work on thecodonts, and in June 1970 I spent
a week in Tucuman studying this material. I found no identifiable
forms not already present in the Harvard-La Plata collections,
BREVIORA
No. 373
but in a number of regards his materials supplemented ours and
substantiated our conclusions. I am deeply grateful to Sr. Bona-
parte and the authorities of the Instituto Lillo for placing this
material at my disposal.
LuPEROsucHUS FRACTUS gen. et sp. nov.
Combined generic and specific diagnosis. A large thecodont,
with an estimated skull length of about 60 cm, probably pertain-
able to the family Rauisuchidae. A shtlike opening apparently
present posterior to the nares between premaxilla and maxilla;
antorbital opening large; apparently no parietal foramen; lateral
temporal opening with vertical posterior border; archosaur type
of otic notch partially developed.
Holotype of the species. La Plata Museum 1964-X1-14-9, an
incomplete skull, consisting of most of the dermal roof and part
of the left side of the "face" collected from the Chanares Forma-
tion in La Rioja Province, Argentina, north of the north fork of
the Chanares River, about 5 km NE of the point where this river
emerges into the Piano de Talampaya.
The generic and specific names refer to the fragmentary and
perplexing nature of the type material.
I am indebted to National Science Foundation Grant GB-2454
for aid in the collecting of the material and to further grants for
its preparation and for publication costs.
Figure 1. Side view of the fragmentary type skull of Liiperosiicluis frac-
tus. as preserved. X 1/6.
197
LUPEROSUCHUS SKULL
Description. This form is represented by a single fragmentary
specimen that includes most of the dorsal surface of a skull and
part of the dermal bones of the left side of the skull (Figs. 1,2).
The condition of the material is none too good, and sutures are
generally difficult to determine. The specimen was found close
beside the skeleton of a dicynodont. Near it were found a con-
siderable number of weathered scraps of bone; whether they
belong to the specimen here described or to the dicynodont
is uncertain.
The animal was a large one; the portions of the skull preserved
measure 54.5 cm in length and if the missing anterior and posterior
regions be restored, the length in life would have been about 60
cm. In general, the reptiles present in the Chanares Formation
are of modest size; apart from this Liiperosuchus specimen, the
dicynodonts are the only large animals known.
As mentioned, most of the skull roof is preserved; the cranium
was obviously long and slender, as in many early archosaurs.
Posteriorly, the parietals are incomplete, and their posterior exten-
sions, which presumably formed the median boundaries of the
superior temporal fenestrae, are missing. There was no parietal
foramen in the portion of the bone preserved (although it may
possibly have been present in the missing posterior portion). A
median longitudinal suture can be made out for almost the entire
length of the roof as preserved. Other sutures are obscured by
poor preservation and bone fusion in this seemingly mature skull.
Figure 2. Dorsal view of the skull, restored. Parts present in stipple.
Abbreviations: /. frontal; /. jugal; /. lacrimal; m, maxilla; /;. nasal; p, pari-
etal; pj, postfrontal; pm. premaxilla; po, postorbital; prj. prefrontal; qj.
quadratojugal; sq, squamosal. X 1/6.
4 BREVIORA No. 373
I have restored the probable sutural pattern, but it should be
emphasized that uncertainties exist.
The area of the posterior part of the frontals and the median
portion of the parietals is depressed, and bounded on either side
by prominent ridges running back along the lateral margins of
the frontals and continuing backward along the parietals. Postor-
bitals are present along the back portion of the supraorbital rims
and extend backward and medially to meet the parietals along
the anterior border of the superior temporal fenestrae. As far as
can be determined, there were large postfrontals, forming part
of the upper margins of the orbits and extending back between
frontals and postorbitals to gain contact with the parietals. The
frontals are broad posteriorly; anteriorly they become reduced in
width between the prefrontals. It is possible that the frontals
entered the orbital margins briefly, but imperfections of the speci-
men render this uncertain, and they may have been excluded by
a narrow contact between postfrontals and prefrontals. The latter
elements appear to be relatively narrow, projecting somewhat
outward over the anterodorsal corner of the orbits and extending
a modest distance forward on either side on the dorsal surface.
Anteriorly, about opposite the front margin of the antorbital vacu-
ity, the nasals, as seen in side view, rise upward markedly above
the general line of the skull roof on their forward course. I was
at first inclined to believe this appearance was due to post-mortem
distortion; however, inspection indicates that this "roman nosed"
elTect is a real structural feature; the conjoined nasals form a
sharp ridge below which the two bones are apposed for some dis-
tance and then slant outward toward either side ventrally. It
seems obvious that the nasals are incomplete anteriorly. Ventrally
the point of separation of the nasal from the posterior extension
of the premaxilla below it indicates the probable position of the
posterior angle of the naris. Most of the premaxilla is missing.
A small fragment of bone attached to the anterior end of the
maxilla presumably represents the most posteroventral position
of the bone. More dorsally a band of bone with well-defined
margins extends dorsoposteriorly between maxilla and nasal, indi-
cating a posterior extension of the premaxilla that excludes the
maxilla from the narial margin, as in certain other thecodonts
(and ornithischians). The whole aspect of the anterior portion
of the skull, as far as preserved, strongly suggests the presence
1971 LUPEROSUCHUS SKULL 5
of an expanded narial area, and in Figure 3 I have restored the
narial region on this supposition.
A large portion of the maxilla is present. A ventroanterior area,
much thickened, represents the ventral margin adjacent to the
premaxilla. This region is excavated internally and, although
preservation is imperfect, represents the area of insertion of a
series of large, probably subthecodont, anterior maxillary teeth.
Above this region the anterior margin of the maxilla slants upward
and backward parallel to the posterodorsal extension of the pre-
maxilla. In the specimen as preserved the two bones are separated
here by a long if narrow slit. For much of this distance the facing
margins of both bones are broadened so that they can readily be
apposed to one another; hence, when I first attempted a restora-
tion of the skull, I placed these margins in firm apposition. But
in contrast to the close union of all other portions of the skull,
in the specimen as preserved, there was here a very distinct separa-
tion, suggesting that a slitlike opening was present in life. Dr.
W. D. Sill, who is currently studying Saurosuchiis, a seemingly
related form from the Ischigualasto Formation, informs me that
such an opening was definitely present in that genus, and I have
therefore indicated such an opening in the restoration in Figure
3. I have no worthwhile suggestion as to the possible function
of this slit.
Back of the nasal region, dorsal and lateral surfaces are sharply
separated for most of the skull length and, even allowing for
possible crushing, it seems certain that the side walls descended
nearly vertically from the lateral dorsal ridges. A short length
of maxilla is preserved ventrally; above, there is a broad plate of
bone apparently formed by the maxilla, extending back above the
antorbital fenestra. The margins of this fenestra are, for the
most part, clearly outlined; it was an opening of considerable size.
In many advanced thecodonts the fenestra is centrally situated in
a depressed area of the cheek; in this specimen the anterior rim
of this depression is clearly incised in the maxilla. The suture
between prefrontal and lacrimal is not clear, but the latter bone
apparently includes the posterior part of the upper margin of the
antorbital fenestra as well as the preserved portion of a stout bar
of bone that separates orbit and antorbital fenestra. Behind the
orbit, the bar of bone between orbit and lateral temporal fenestra
is completely preserved, and there are indications of a suture well
BREVIORA
No. 373
down this bar, between postorbital and jugal. A fraction of the
latter bone is present, defining the lower margin of the orbit, a
section of the cheek rim, and a small area of the anteroventral
margin of the lateral temporal fenestra. Above this fenestra a
stout bar of bone is present, presumably formed anteriorly by
the postorbital, posteriorly by the squamosal (the suture between
the two is not clear). An incomplete flange of the latter bone
extends directly downward as part of the posterior border of the
lateral fenestra. The squamosal extended backward beyond the
level of this descending flange, although this extension is broken
off in the specimen. The vertical descent of the squamosal flange
indicates that the posterior border of the fenestra had not acquired
the V-shaped contour seen in various more advanced thecodonts;
on the other hand, the posterior prong of the squamosal suggests
the initiation of a typical archosaur type of otic notch.
In Figure 3 I have freely restored the skull in side view to give
a suggestion of its probable appearance in life. Despite the incom-
plete nature of the evidence I do not think that there can be too
great a departure from life conditions in most regards. Most
doubtful, because of lack of material, is the suspensorial region.
Systematic position. As to relationships of Luperosuchiis, an
early Middle Triassic form, one tends to think first of the larger
erythrosuchid members of the Proterosuchia — a group most
recently discussed by Reig (1970), and by Charig and Reig
(1970). Primitive, for example, is the apparent presence of a
large postfrontal. Liiperosuchus, however, is more advanced than
Figure 3. Side view of the skull, restored. Abbreviations as in Fig.
2. X 1/6.
1971
LUPEROSUCHUS SKULL
proterosuchians in various regards, such as the large size and
incised nature of the antorbital fenestra, probable absence of a
parietal foramen, and the apparent beginning of the pseudosuchian
development of an otic notch. More reasonable is assignment to
the Rauisuchidae (or Prestosuchidae), a family of large but rela-
tively primitive Middle Triassic thecodonts, first seriously studied
by Reig (1961). Included here may be such forms as Ticinosii-
chiis from the European Anisian (Krebs, 1965), Fenhosuchus
and, doubtfully, Shansisiichus from China (Young, 1964), Stcigo-
nosuchus (Huene, 1938), and Mandasuchus from the Manda
beds of East Africa. The presence of rauisuchids in the Middle
Triassic of South America is well attested by the presence of
Rauisiichiis and Prestosuclms from the Santa Maria of Brasil and
Saiirosuchus of the Middle Triassic Ischigualasto Formation of
Argentina.
The material of Liiperosuchus is too fragmentary to warrant
any extended discussion of rauisuchid relationships. Are they, as
Reig believes (1970, fig. 10), a side branch from a somewhat
advanced pseudosuchian stock, or could they have progressed in
parallel fashion from the proterosuchian base of the Thecodontia?
Are they a sterile group, without descendants, or could they be
related to the ancestry of certain of the later saurischians, the
Prosauropoda (Palaeopoda) or, more especially, ancestral Sauro-
poda? It is possible that Dr. SilFs current studies of Saurosuchus
will shed light on rauisuchid relationships.
Dermal scutes. In two instances we found in the Chanares
region large scutes not definitely associated with other identifiable
skeletal remains (Fig. 4). They are too large to be attributed to
any of the other (and much smaller) thecodonts present in our
1 2 CM
J I
Figure 4. Two dermal scutes, possibly referable to Luperosuchus.
8 BREVIORA No. 373
collections (and they are not, of course, attributable to the synap-
sids, which make up the remainder of the materials collected).
Of known forms from the Chanares, Luperosuchus is the only
one to which they could have belonged and, since comparable
scutes are known in other rauisuchids, we may provisionally assign
them to the present genus. None of the scutes is perfectly pre-
served. One type, rectangular in shape, has a thickened, saw-
toothed border along one edge, indicative of an interdigitating
connection with another element. Such scutes are presumably
paramedian paired scutes, found in various other thecodonts.
Subcircular scutes, also present, may be more lateral elements or
median caudal ones.
REFERENCES CITED
Charig, a. J., AND O. A. Reig. 1970. The classification of the Protero-
siichia. Biol. Jour. Linn. Soc, 2(2): 125-171.
HuENE, F. VON. 1938. Ein grosser Stagonolepide aus der jiingeren Trias
Ostafrikas. Neues Jahrb. Min. Geol. Pa!., Beii.-Bd., 80: 264-278.
Krebs, B. 1965. Ticinosuchiis ferox nov. gen. nov. sp. Ein neuer Pseudo-
suchier aus der Trias des Monte San Giorgio. Schweiz. Palaont. Ab-
handl., 81: 1-140.
Reig, O. A. 1961. Acerca de la Posicion Sistematica de la Familia
Rauisuchidae y del Genero Saiirosiichiis (Reptilia, Thecodontia).
Publ. Mus. Munic. Cienc. Nat. Tradic. Mar del Plata, 1(3): 73-114.
1970. The Proterosuchia and the early evolution of the
archosaurs; an essay about the origin of a major taxon. Bull. Mus.
Comp. Zool., 139(5): 229-292.
Young, C. C. 1964. The pseudosuchians in China. Palaeont. Sinica, new
sen C, No. 19: 107-205.
BREVIORA
Mmseiuijni of Comparative Zoology
Cambridge, Mass. 15 June, 1971 Number 374
THE FISHES OF THE MALAYSIAN FAMILY
PHALLOSTETHIDAE (ATHERINIFORMES)
Tyson R. Roberts^
Abstract. The tiny fishes of the family Phallostethidae, from Malaya
and Thailand, develop a large penis and differ radically in structure of the
bilaterally asymmetrical priapium from the somewhat larger fishes of the
more widely distributed Neostethidae, the only other family in the sub-
order Phallostethoidea. A morphological characterization of the Phallo-
stethidae is given and its features compared to those of Neostethidae.
There are three species: Phallostethus diinckeri Regan (1913), known
only from the type specimens collected in the mouth of the Muar River
in Johore, Malaya; Phenacostethus smithi Myers (1928), known from the
types and many other specimens collected in the khlongs of Bangkok and
reported in this paper from Chantaburi Province in southeast Thailand;
and Phenacostethus posthon, new species, from the Indian Ocean coast of
peninsular Thailand. Ph. posthon and Ph. smithi differ considerably in
morphology of the priapium and penis. Furthermore, the asymmetrical
priapium in Ph. posthon is invariably sinistral. In all other phallostethoids,
so far as known, it may be either sinistral or dextral; in Ph. smithi the ratio
of sinistral and dextral males is near equality (Hubbs and Hubbs, 1945).
The ecology of phallostethids is described for the first time, excepting
some brief remarks by H. M. Smith (1927; 1945), who seems to have
confused Phenacostethus in the field with neostethids and perhaps with
Oryzias. An hypothesis is offered that the selective advantage of internal
fertilization in Phallostethoidea (an oviparous group) lies in permitting
temporal separation of mating and spawning activities, corresponding, re-
spectively, with periods of low water and high water in habitats subject
to strong tidal fluctuations. Two trends in the reproductive biology of
atheriniform fishes that might be conducive to the evolution of internal
fertilization are: 1) towards eggs in which either embryonic development
is slowed down or temporarily arrested, or hatching of embryos is deferred;
and 2) away from expelling all ovulated eggs at once and towards expel-
ling them in small batches or even singly.
1 Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts 02138.
2 BREVIORA No. 374
ACKNOWLEDGMENTS
My studies in Thailand were carried out under the sponsorship
of the Thai National Research Council. The College of Fisheries
of Kasetsart University acted as my host, providing laboratory
space, library facilities, and access to fish collections, including
specimens of Phenacostethus smithi collected by H. M. Smith. I
wish to thank the following people for their substantial aid, par-
ticularly in the aspects of my fieldwork concerning phallostethoids:
Dean Jinda Thiemeedh, Mr. Prajit Wongrat, and Mrs. Supap
Monkolprasit of the College of Fisheries; Miss Prachuab Suk-
charean and Mr. Sopon Chantarat of the Marine Fisheries Station
of Songkhla; and Dr. Vagn Hansen, Director of the Phuket Marine
Biological Center. Mrs. Monkolprasit was extremely helpful, par-
ticularly in arranging my trips. Dean Thiemeedh kindly en-
couraged my work and arranged transportation to Chantaburi
Province, where, with Mr. Wongrat's help, not only Ph. smithi,
but also large series of the neostethids Neostethus siamensis
(hitherto known only from a single female) and Ceratostethus
bicornis (previously unrecorded from Thailand) were obtained.
Mr. Wongrat also helped find Phenacostethus near Bangkok.
Miss Sukcharean arranged my travels from Songkhla to Satul; Mr.
Chantarat accompanied me on this trip and helped collect the
first specimens of Phenacostethus posthon. Dr. Hansen arranged
my fieldwork in Pungah.
For translations of the papers by Aurich and Woltereck I am
obliged to Miss Deborah White and Dr. Elizabeth Deichmann.
Prof. George S. Myers read the paper in manuscript.
INTRODUCTION
This account of the family Phallostethidae is the first paper
dealing with the fresh- and brackish-water fishes collected by me
in Thailand from April 15 to July 14, 1970, and deposited in the
fish collection of the Museum of Comparative Zoology. Collecting
phallostethoid fishes was one of the main objectives of my field-
work in Thailand. In addition to Phenacostethus smithi Myers
(1928) and the new phallostethid described in this paper, large
series of the neostethids Neostethus siamensis Myers (1937)
and Ceratostethus bicornis (Regan, 1916) were obtained. These
represent the first specimens of Ceratostethus recorded from Thai-
land, and the only specimens of Neostethus siamensis other than
1971 PHALLOSTETHIDAE 3
the female holotype. (/V. siamensis is close to, and perhaps spe-
cifically identical with, N. lankesteri Regan (1916), the type
locality of which is the mouth of the Muar River, Johore, and
Singapore.) The osteology and functional anatomy of Cerato-
stethus will be considered in another paper.
The Phallostethoidea are small, highly specialized fresh- or
brackish-water fishes in which males have a remarkable sub-
cephalic copulatory organ, the priapium. The skeleton and mus-
culature of this complicated bilaterally asymmetrical organ, which
functions both in clasping and intromission, are derived mainly
from the pelvic fins and girdle. Minor contributions come from
the first pair of ribs and anteroventral part of the pectoral girdle.
In females the pelvic fins are absent or vestigial. Aurich (1937)
divided the Phallostethoidea into two "Familien," but gave these
divisions names in subfamily form, Phallostethinae and Neo-
stethinae. Berg (1940: 465-466) recognized them as families,
Phallostethidae and Neostethidae, as did Rosen (1964: 261)
and Greenwood et al. (1966: 398). Neostethidae, comprising
eight genera and about 15 species, have been recorded from
Thailand, Malaya, Sumatra, Borneo, and the Philippine Islands.
Six of the genera — Gulaphallus Herre (1925), Mirophallus
Herre (1926), Plectrostethiis Myers (1935), Solenophallus Au-
rich (1937), Ctenophallus Herre (1939), and Manacopus Herre
(1940) — are known only from the Philippines.
The Phallostethidae, even more specialized (and rarer in mu-
seum collections) than Neostethidae, comprise three species from
the Malay Peninsula and adjacent parts of Thailand. Phallostethus
dunckeri Regan (1913), the first phallostethoid to be described,
is known only from the type specimens Duncker collected previous
to 1904 at the mouth of the Muar River, about 20 miles south
of Malacca, Johore Province, Malaya (Duncker, 1904: 171).
Phenacostethus smithi, hitherto known only from several large
series collected by H. M. Smith from khlongs in Bangkok, was
obtained by me at Bangkhen (a suburb of Bangkok) and at
Chantaburi, near the southeast corner of Thailand (near Cam-
bodia). The third species, described in this paper, is from the
Indian Ocean coast of Thailand. All localities where phallostethids
have been collected are shown in Figure 1. The present paper
presents characterizations of the family Phallostethidae and of
the phallostethid species, a description of the new species just
mentioned, and observations on the ecology of Phallostethidae.
BREVIORA
No. 374
ir
10"
5*-
- 10"
15*
5*
100*
lOS*
Figure 1. All localities where Phallostethidae have been collected. 1.
Mouth of Muar River at Bandar Maharani (type locality of Phallostethus
dimckeri Regan, 1913); 2. Bangkok (type locality of Phenacostethiis smithi
Myers, 1928); 3. Bangkhen, a suburb of Bangkok (Ph. smithi); 4. Chanta-
buri City {Ph. smithi); 5. Khlong La Ngoo, 48 km NW of Satul Town,
6° 52' 30" N, 99° 48' 10" E (type locality of Phenacostethiis posthon n.
sp.); 6. Khlong Kla Sohm, 15 km S of Pungah Town, on Pakasem Road
from Pungah going towards Phuket Island (type locality of Ph. posthon
n. sp.)
Definition of the suborder Phallostethoidea and discussion of
phyletic trends in the Phallostethoidea will be presented in a
forthcoming paper on Ceratostethus.
Ph. smithi provides an Asian example of the phenomenon (of
which Africa and South America provide numerous examples)
1971 PHALLOSTETHIDAE 5
that the smallest fishes amidst the richest tropical freshwater
faunas are representatives not of the dominant fish groups present,
but of groups with a marginal distribution pattern. The ecology
of such minute fishes, when known, usually proves to be highly
specialized.
Rosen (1964) united the exocoetoids, scomberesocoids, adri-
anichthyoids, cyprinodontoids, atherinoids and phallostethoids in
a new order, the Atheriniformes. Atherinoids and cyprinodont-
oids were widely separated in earlier classifications, the artificial-
ity of which has become increasingly apparent. Such similarities
as were noted between atherinoids and cyprinodontoids had
usually been attributed to convergence. Having personally investi-
gated the osteology of phallostethoids (which evidently are re-
lated to atherinoids), atherinoids, and cyprinodontoids, and
reviewed much of the literature on osteology and reproductive
biology of these groups, I am inclined to believe that they may
be related. Some of the similarities in reproductive biology were
first brought to my attention in a talk given by Neal R. Foster
at the 1968 meetings of the American Society of Ichthyologists
and Herpetologists.
Note on the figures. Figures 2-5, prepared with the aid of a
Wild microscope and camera lucida, are based on formalin speci-
mens. Formalin specimens of Phallostethidae are more nearly
normal in appearance than alcoholic specimens, since the latter
invariably undergo at least some shrinkage. In specimens that
have been transferred from the original formalin fixative to 60
per cent ethyl alcohol for permanent storage, teeth protrude more
from the gums, scales stand out more clearly, the membranous
dome atop the head has disappeared, and the caudal peduncle is
decidedly narrower.
CHARACTERIZATION OF THE FAMILY PHALLOSTETHIDAE
The following characterization of the Phallostethidae is based
on my observations of Ph. sinithi and Ph. posthon, including study
of alizarin preparations, and on accounts of Ph. smithi by Myers
(1928), Bailey (1936), and TeWinkel (1939), and of Phallo-
stethus dunckeri by Regan (1913; 1916).
1 . Slender elongate phallostethoids, very delicate, largely trans-
lucent, with deciduous scales; externally visible concentrations of
melanophores restricted to the top of the braincase, middle of the
6 BREVIORA No. 374
dorsum, midlateral intermuscular septum, priapium, and bases and
edges of fin rays; maximum standard length about 23 or 24 mm.
2. Dorsum of head with a translucent, membranous dome.
3. Mature individuals of both sexes with a bright orange-yellow
bar on caudal peduncle.
4. Branchiostegal rays 4.
5. Main "externalized" clasping bone in the priapium is the
toxactinium; ctenactinium reduced or absent.
6. A greatly enlarged, oval, concave pad, or pulvinulus, shghtly
posterior to toxactinium.
7. Vas deferens terminating in a large penis that projects con-
siderably from the priapium.
8. Pelvic spines or rays, if present, greatly reduced and modi-
fied beyond recognition.
9. Vas deferens highly coiled, forming a sort of epididymis.
Comments on the family characters. Neostethids, while small
as fishes go, are all or almost all larger when adult than phallo-
stethids. Most, if not all, neostethids are hardier fishes than phal-
lostethids and have relatively adherent scales. While they also
are largely translucent, neostethids usually have relatively more
melanophores than phallostethids. The epidermis paralleling the
scale margins is often well provided with melanophores in neo-
stethids but invariably devoid of melanophores in phallostethids.
Neostethids (Ceratostethus and Neostethus), even at comparable
sizes, lack a membranous dome on the dorsum of the head, or if
one is present, it is not notably elevated. According to TeWinkel
(1939) this region bears sensory canals in both Phenacostethus
and Gulaphallus. In neostethids large sensory pores are evident
in the frontal region, whereas the membranous dome in phal-
lostethids is apparently entire.
With regard to characteristic number 3 (bright orange-yellow
markings at the base of the caudal fin and sometimes at the origin
of the anal fin), these are present in Uving specimens of Ph. smithi
and in Ph. posthon. There is no way of teUing whether such spots
are also present in Phallostethus. The orangish yellow coloration,
contained in chromatophores (approximately 50-75 chromato-
phores constituting the caudal base mark) gradually disappeared
after a few weeks of preservation in formalin. Similar markings
1971 PHALLOSTETHIDAE 7
were definitely absent in live specimens of Neostethus and Cera-
tostethus observed by me, nor is there any mention of such marks
in the literature on Neostethidae.
Concerning character 4, Neostethidae usually have 5 branchio-
stegal rays. The number of branchiostegal rays in Phallostethus
is unknown.
Characters 5-9 concern the priapium, which differs funda-
mentally from that of Neostethidae. In Neostethidae the main
"externalized" bony clasping element, the ctenactinium, is appar-
ently a modified pelvic fin ray or spine. The mam "externalized"
bony element in the priapium of phallostethids, the toxactinium,
is not homologous with the ctenactinium. My observations con-
firm Bailey's view (Bailey, 1936: 463, 471) that it is homologous
with the pulvinular bone, one of the anteriormost internal bony
elements in the priapium of neostethids. The homologies of this
element are unclear, but it is almost certainly not a modified pel-
vic ray or spine. The ctenactinia of phallostethids, which may or
may not be homologous with the elements called ctenactinia in
neostethids, are greatly reduced in size. The ctenactinium of Phal-
lostethus dimckeri, while relatively short, bears several "teeth"
or sharp projections; these are perhaps comparable to the single
curved hook present near the base of the ctenactinium in Neo-
stethus. The pulvinulus of phallostethids probably functions as a
pad in conjunction with the toxactinium. A homologous but much
smaller pulvinulus is present in neostethids. A striking character-
istic of the phallostethid priapium is the development of a large
penis. In Phenacostethus posthon the organ is entirely smooth;
in Ph. smithi its distal half bears a series of stiff ruffled pleats.
The development of a large penis evidently occurred independently
in the neostethid Mirophallus bikolanus (Herre, 1926, pi. 3, fig.
1 ) . In most neostethids a compUcated flap covers the opening of
the vas deferens (Aurich, 1937). This flap is absent in PhaUo-
stethidae. Various bony elements in the priapium of phallostethids,
including a peculiar slender element lodged in the concave side
of the penis bone in Ph. smithi, may be homologous with pelvic
rays, but the priapium of phaUostethids does not bear any
branched elements that obviously are relatively unmodified pelvic
rays. Several branched pelvic rays of relatively normal appear-
ance occur in the neostethids Ceratostethus, Neostethus, Soleno-
phallus, and Gulaphallus (personal observations; Aurich [1937],
TeWinkel [1939], Woltereck [1942 a, b]).
8 BREVIORA No. 374
The priapium of Phallostethus is clearly of the same general
type as that of Ph. smithi and Ph. posthon. In Phallostethus the
toxactinium and pulvinulus are very similar to these structures in
Phenacostethus. It is likely that Phallostethus develops a large
penis. The penis in phallostethids, like the ctenactinium of neo-
stethids, only reaches its full development in the largest males.
Regan (1916: 22) hypothesized that in Phallostethus the tox-
actinium grips the female under the chin or is held in her mouth,
while the serrated edge of the ctenactinium gives a firm hold on
the pectoral region in front of and on the far side of the genital
orifice, in order that the seminal papilla could be placed against
it or introduced into it. In Neostethus he hypothesized that the
female is held across the back of the head by the ctenactinium,
the anterior descending part of which lies on the side of the female
away from the male. Copulation has yet to be observed in Neo-
stethus or in any phallostethids. It has been observed only in the
neostethid Gulaphallus mirabilis. In this species the female is
held across the back of the head by the ctenactinium; the "second
ctenactinium" of Gulaphallus mirabilis, which actually is an exter-
nalized pelvic bone, apparently rests or presses against the female's
opercular region on the side next to the male (Villadohd and Man-
acop, 1934: pi. 5, fig. 2). The reduced ctenactinium of Phallo-
stethidae could hardly function in the same manner as the elongate
ctenactinium characteristic of all neostethids. It may be that the
toxactinium is held, not under the female's chin or in her mouth,
but atop the front of her head.
In Phallostethus (Regan, 1916: 19, fig. 14) the vas deferens
is highly coiled within the abdominal cavity to form a sort of
epididymis. TeWinkel (1939) reports a similar coiling of the
vas deferens in Ph. smithi. In Neostethus lankesteri (Regan,
1916: 10, fig. 6) and in Gulaphallus mirabilis (Villadolid and
Manacop, 1934: pi. 3, fig. 4) the vas deferens is unconvoluted
from its origin on the testis to where it enters the priapium, then
forms an expanded loop inside the priapium. The sperm of Neo-
stethus lankesteri (Regan, 1916: 13, fig. 9) are concentrated into
"spermatophores," more properly called spermozeugmata (Niel-
sen et al., 1968: 248). Regan (1916: 19) stated that Phal-
lostethus evidently did not produce "spermatophores" like those
of Neostethus. In mature Neostethus and Ceratostethus (per-
sonal observation) the posteriormost portion of the priapium is
1971 PHALLOSTETHIDAE 9
sometimes greatly swollen with closely packed, adherent small
vesicles that presumably are spermozeugmata. The priapium
apparently does not become similarly swollen in phallostethids.
Insofar as can be determined from examination of the bones
only, the contributions of the first pair of ribs and of the
shoulder girdle to the priapium is the same in Phallostethidae and
Neostethidae.
THE SPECIES OF PHALLOSTETHIDAE
Phallostethus dunckeri Regan 1913
Phallostethus dunckeri Regan, 1913: 550, figs. 1-4 (original description;
types from mouth of Muar River at Bandar Maharani, Johore, Ma-
laya; soft anatomy). — Regan, 1916 (soft anatomy, histology, oste-
ology, comparison with Neostethiis).
This species is known only from the specimens collected by
G. Duncker at the mouth of the Muar River at Bandar Maharani,
Johore, Malaya. The following statements are based on the two
accounts of this species by Regan (1913, 1916). It is character-
ized by an exceptionally long anal fin, with 26-28 elements
(eight to ten rays more than are found in phallostethoids with
the next highest number of anal fin elements), an anal base about
30 per cent of the standard length (compared to anal base 20-25
per cent of standard length in all other phallostethoids) and origin
of anal considerably nearer to snout tip than to end of hypural
fan (anal origin slightly to considerably nearer to end of hypural
fan than to snout tip in all other phallostethoids), and by a ser-
rated ctenactinium (Regan, 1916: fig. 13). Regan did not have
very many specimens (some were used for histological preparations
or cleared in oil of cloves for bone study). Dr. P. H. Greenwood
informs me that the British Museum (Natural History) has four
specimens (types?) of Phallostethus dunckeri in rather poor con-
dition. Although Regan's specimens of Phallostethus dunckeri
(at 23-29 mm in total length) are larger than Phenacostethus,
and the males figured by him have well-developed priapia, perhaps
the penis is not fully developed in them. This seems likely, be-
cause in both species of Phenacostethus the largest females are
only about one-half to one milUmeter longer than the largest
males, whereas Regan's largest female of Phallostethus is six mil-
limeters longer than his largest male. In Phenacostethus the penis
is fully formed only in the largest males.
10 BREVIORA No. 374
The second dorsal fin of P. dimckeri has eight to ten elements
(a number found in some Neostethidae); the highest number in
Phenacostethus is seven. It is unknown whether Phallostethus has
a first dorsal fin. Regan did not mention its presence, but he over-
looked the first dorsal in his material of Neostethus lankesteri
and Ceratostethus bicornis. Phallostethus probably has a small
spinous first dorsal fin, but the absence of one would not be unique
among Phallostethoidea: the first dorsal is lacking in the neosteth-
ids Mirophallus bikolamis Herre (1926), (Herre, 1942: 141;
Myers, 1937: 142) and Solenophallus thessa Aurich (1937: 265).
Phenacostethus smithi Myers 1928
(Figures 2, 4, 6)
Neostethus lankesteri (not of Regan) Smith, 1927: 353-355 (misidenti-
fication).
Phenacostethus smithi Myers, 1928: 6, figs. 1 and 2 (original description;
types from Bangkok; holotype in American Museum of Natural His-
tory). — Bailey, 1936 (osteology).— TeWinkel, 1939 (soft anat-
omy).—Smith, 1945: 475 (Bangkok; synonymizes Ph. thai Fowler;
ecology).
Phenacostethus thai Fowler, 1937: 219; figs. 189, 190 (original description;
Bangkok; holotype in Philadelphia Academy).
Material studied. MCZ 47055, 13 specimens, five females
13.7-14.8 mm and eight males 13.6-14.9 mm (three with toxac-
tinium arising on left side, five on right side), from Khlong Bang-
khen at bridge on Nzarm Wong Wan Road, a few km west of
Kasetsart University, Bangkhen, Bangkok, Thailand, 30 April
1970; MCZ 47299, 20 specimens, three females 13.4-15.6 mm
and 17 males 12.7-14.9 mm (12 with toxactinium arising on right
side, five on left side), from Khlong Kee Nawn, behind Cathohc
church in Chantaburi City, Chantaburi Province, Thailand, 5
May 1970.
Myers' figure 1 of Ph. smithi does not show the membranous
dome of the dorsum of the head (Fig. 2) ; it is likely to be shrunken
in alcoholic specimens. The adult male in Myers' figures 1 and 2,
at 13.5 mm in standard length, does not have the penis as fully
developed as in my specimens 14.3 (Fig. 2), 14.1 (Fig. 4), and
14.5 (Fig. 6) mm in standard length. In these specimens the
penis is much larger, and its distal end bears a series of a half-
dozen or more crenulated radial folds or extensions resembling a
1971
PHALLOSTETHIDAE
11
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12 BREVIORA No. 374
set of ruffled lace cuffs (best shown in Fig. 4). These folds can
be seen in one of Bailey's anatomical figures based on material
of Ph. smithi collected by Hugh M. Smith. Direct comparison of
Ph. smithi collected by Smith in Bangkok (deposited in the fish
collection of the College of Fisheries, Kasetsart University, and
in the MCZ fish collections) with my material from Bangkhen and
Chantaburi indicates that only one species is involved. Live speci-
mens of Ph. smithi of both sexes from Bangkhen and from Chanta-
buri had a bright orange-yellow, anteroventrally-posterodorsally
oriented bar on the lower half of the caudal peduncle. This bar
disappeared from specimens kept a few weeks in formahn. The
color was due to about 50-75 chromatophores. There is a small,
shiny blue spot over the brain (well behind the eyes), and a very
small silvery spot in the middle of the eyes' dorsal surface.
Alizarin preparations reveal the first dorsal fin consists of a
single tiny spine, as indicated by Myers. Most specimens are miss-
ing many scales. Head scaleless. Abdominal keel scaleless in
both sexes (abdominal keel scaled in females of Ceratostethus
bicornis). Scales on body cycloid, number of scale rows corre-
sponding closely with the number of myotomes. Teeth in both
jaws in a single series; medial portion of premaxillary with about
eight sharp conical teeth, lateral (expanded) margin of premaxil-
lary with about seven to nine slightly larger conical teeth; medial
portion of dentary with about a dozen conical teeth opposing but
much smaller than those on medial portion of premaxillary.
For counts of fin elements and vertebrae based on alizarin
preparation see Table 1.
Phenacostethus posthon, new species
(Figures 3, 5, 7)
Holotype. MCZ 47300, a 16.7-mm male from Khlong Kla Sohm
about 15 km southwest of Pungah Town, where it is crossed by
a bridge on the Pakasem Road (between Pungah Town and Phu-
ket), Pungah, Thailand. 29 June 1970.
Paratypes. MCZ 47301, 58 specimens, comprising two im-
matures, 11.9 and 12.0 mm, 22 females, 12.3-17.7 mm, and 34
males 12.7-17.0 mm (of which two males, 15.5 and 16.5 mm, are
cleared and stained), same data as holotype; MCZ 47302, 20
specimens, comprising three immatures 9.6-12.0 mm, 10 females
12.7-18.0 mm, and 7 males 12.9-16.7 mm, from Khlong Langu
1971 PHALLOSTETHIDAE 13
at Langu Town, 48 km northwest of Satul Town (6° 52' 30" N,
99° 48^' 10^' E). 23 June 1970.
The most obvious differences between this species, from the
Indian Ocean coast of Thailand (Fig. 1), and Ph. smithi involve
the priapium and the position of the dorsal fin. Granted that both
species have the characteristic priapial elements of the family
Phallostethidae, the priapium is, in fact, so different in the two
species that the question may even be raised as to whether separate
generic status is indicated.
Differences in the priapium of the two species involve external
morphology of the penis, skeleton of the penis, ctenactinium, tox-
actinium, and the nature of the laterality of the priapium itself.
In Ph. smithi 1 ) the distal portion of the penis is ruflfled (see
description above); 2) the penial skeleton includes a large papil-
lary bone with a slender penial bone lodged in its concave surface
(Fig. 6; see also Bailey, 1936: 3 and 4); 3) the ctenactinium is
relatively large and externally evident; 4) the toxactinium is rela-
tively slender and gently curved; and 5) the priapium itself may
be either sinistral or dextral. In Ph. posthon, on the other hand,
1 ) the penis is smooth; 2) the penial skeleton has a large papillary
bone but the penial bone is absent; 3) the ctenactinium, if it is
present at all, is reduced and hardly detectable externally; 4) the
toxactinium is stouter and distinctly more sharply curved; and 5)
the priapium is invariably sinistral (toxactinium arising on left
side) in the material examined.
Ph. posthon and Ph. smithi of both sexes can be distinguished at
a glance by the position of the first dorsal fin relative to the anal
fin base. In Ph. posthon the first dorsal (which has a single spine
as in Ph. smithi) originates slightly posterior to the base of the
last ray in the anal fin; in Ph. smithi it originates over the middle
of the anal fin base. The first dorsal fin is somewhat closer to
the second dorsal origin in Ph. posthon than in Ph. smithi. Ph.
posthon is more elongate (depth of body at anal fin origin about 7,
vs. 6 in Ph. smithi) and evidently a larger species. The average
and maximum sizes of specimens in my two samples of Ph.
posthon are definitely larger than in my two samples of Ph. smithi
(which include specimens larger than those previously recorded).
The largest male and female specimens of Ph. smithi are, respec-
tively, 14.9 and 15.6 mm; of Ph. posthon, 17.0 and 18.0 mm
(standard lengths). The largest specimens (both sexes) had a
bright orange-yellow bar on the caudal peduncle, as in Ph. smithi,
14
BREVIORA
No. 374
Figure 4. Phenacostethus smithi Myers (1928), ventral view of pri-
apium, 14.1-mm specimen, MCZ 47055 (formalin).
and also a smaller orange-yellow bar on the body next to the anal
fin origin.
As in Ph. smithi, there appears to be a one-to-one correspon-
dence between the scale rows and the myotomes. The thin scales
are extremely difficult to see in specimens under alcohol or water;
and most specimens are missing many scales. I find in several
specimens eight oblique scale rows between dorsal midline and
anal fin origin, and six oblique scale rows between dorsal fin
origin and anal base. Shape of jaw bones and disposition of teeth
as in Ph. smithi.
For fin and vertebral counts based on alizarin preparations of
Ph. posthon see Table 1. Ph. posthon seems to have, on the aver-
age, one less ray in the dorsal fin (verified by counts of unstained
specimens) and one more vertebra than Ph. smithi.
Right- and left-handedness in the priapiiim of Phallostethidae.
1971
PHALLOSTETHIDAE
15
Figure 5. Ph. posthon n. sp., ventral view of priapium in 16.9-mm para-
type, MCZ 47301 (formalin).
That the priapium of Neostethus lankesteri may be either right-
or left-handed is stated by Regan (1916: 5). Regan did not spe-
cifically state that the priapium of Phallostethus may be either
way, but in a general discussion of the priapium as compared to
copulatory organs of other fishes, he refers to its being either
dextral or sinistral (Regan, 1916: 23). Regan's figures of Phal-
lostethus depict both right-handed and left-handed males. It is
conceivable that one or more of them might be reversed images,
the images being reversed either by the illustrator or by the printer,
so the figures cannot be taken as definite proof that P. dimckeri
may be either left-handed or right-handed. Concerning Phena-
costethus smithi, Hubbs and Hubbs (1945: 294) found that in
334 males (from material collected by H. M. Smith at Bangkok)
16
BREVIORA
No. 374
posttemporal
c lei thrum
coracoid
supracleithrum
scapula
tozactiiiium
antepleural
papillary
axial
Figure 6. Ph. smithi Myers (1928), skeleton of priapium and pectoral
fin in 14.5-nim specimen, MCZ 47301. Terminology of priapial elements
according to Bailey (1936).
posttemporal
cleithrum
coracoid
supracleithrum
scapula
radial
toxactinium
pulvinulus
antepleural
ribs
anal
basipenial
prepapillary
uncus
anterior infrasulcar'
posterior infrasulcar
ctenactinium
Figure 7. Ph. posthon n. sp., skeleton of priapium and pectoral fin in
16.5-mm paratype, MCZ 14301. Priapial elements of uncertain homology
with those in Ph. smithi are unlabelled.
1971 PHALLOSTETHIDAE 17
the aproctal side was the left side in 155 and the right side in
179. The deviation from equality is statistically insignificant. In
my samples from Bangkhen and from Chantaburi (too small for
statistical analysis) I find both left- and right-handed males. In
Phenacostethus posthon the toxactinium arises on the left side
in every male in which the priapium is sufficiently developed to
have a toxactinium (27 specimens from Pungah and five from
Satul). To my knowledge this is the only phallostethoid in which
the sidedness of the priapium appears to be fixed. Females of Phal-
lostethidae are bilaterally symmetrical, the genital openings lying
in the middle of the throat.
ECOLOGICAL OBSERVATIONS
The only ecological information Regan had about Phallostethus
dunckeri was that it came from brackish water from the Muar
River at Bandar Maharani. Duncker also collected Neostethus
lankesteri in the same general locahty.
Smith (1927; 1945: 476) made brief ecological observations
on Phenacostethus smithi but in restrospect it seems that he some-
times confused this little fish in the field with Neostethus or Cera-
tostethus and perhaps even with Oryzias. I did not observe either
phallostethids or neostethids at the surface, and feel that they
generally keep well below the surface. Smith's remarks (that
Phenacostethus "normally remain at or close to the surface, where
they would be difficult to see were it not for a glistening yellow
area on the top of the head'') sound more like Oryzias. The glisten-
ing area on the top of the head in phallostethoids is relatively small
and bluish, compared to the large yellowish glistening area in
Oryzias, which is comparable to the striking head spots in rivuline
cyprinodontids of the genera Epiplatys and Aplocheilus. In any
event. Smith records that Ph. smithi abounds in freshwater pools,
ditches and smaller canals in the Bangkok region, Hving in water
that is nearly always muddy or turbid, and that the species is
oviparous (although spawning was not observed); the egg-bearing
and spawning periods are protracted, corresponding with the rainy
season, and may extend from May to December. Small numbers
were maintained in aquaria for a month by the daily introduction
of fresh ditch or canal water to provide food in the form of minute
crustaceans, protozoans, worms, etc. Smith found the larvae of
anopheline mosquitoes much too large for Ph. smithi to ingest.
18 BREVIORA No. 374
Undoubtedly Ph. smithi once lived in innumerable khlongs and
ditches in Bangkok which are today so polluted that they are
inhabited only by the hardiest air-breathing fishes such as Anabas
testudineiis. In Khlong Bangkhen, a few kilometers from Kasetsart
University, I found Ph. smithi in association with a variety of
primary and secondary freshwater fishes, principal among which
were cyprinids mainly of the genus Rasbora, Dermogenys, Xenen-
todon, Oryzias, Gobiopterus chuno (a small translucent goby),
young Fhita, and Chaudhuria. The Phenacostethus were mostly
in a little backwater, close to the main current in the khlong, in
very turbid water. I was unable to taste any salt in the water.
At Chantaburi City, Phenacostethus was collected in a turbid
ditch or small khlong, Khlong Kee Nawn, behind the Catholic
church. There were a number of houses along the khlong at this
point, and considerable rubbish had been thrown into it, so col-
lecting efl'orts were confined to dip-netting for Phenacostethus.
Dermogenys, Brachygobius and Gobiopterus were collected inci-
dentally. About two kilometers downstream, where the khlong
traversed a large open field, we found minnows such as Oxygaster,
Esomus and Rasbora (but no Phenacostethus) which indicates the
water in the khlong was not brackish.
At Satul and at Pungah Ph. posthon was obtained along the
margins of khlongs or tiny branches of khlongs in which the water
was swiftly flowing, highly turbid, and probably completely fresh.
A few of the largest females (collected June 1970) were replete
with ripe ova. Both localities were at places fully subject to tidal
fluctuations, but far upstream from branches having water the
least bit sahy to the taste. At Khlong Langu in Satul Province
the water level was undoubtedly high because of heavy rains for the
preceding 12 hours or more; Phenacostethus were collected in
the khlong both on the falling tide and rising tide. No primary
freshwater fishes were obtained. Species in the khlong where
Ph. posthon was collected included Oryzias, Dermogenys, Chanda,
Gobiopterus, Butis, and Tetraodon. In Khlong Kla Sohm, near
Pungah, Ph. posthon was coUected in a narrow side channel more
or less uniformly 3-4 feet deep, swiftly flowing, with hard-packed
mud bottom, well shaded by Nipa palm and mangrove. Oryzias,
Dermogenys, Gobiopterus, Periophthalmus, Tetraodon, and young
Scatophagus were the only other fishes collected or observed in
this channel.
1971 PHALLOSTETHIDAE 19
In contrast, on the more numerous occasions when I obtained
either Neostethus or Ceratostethus in Thailand, the water was
usually brackish or even very salty to the taste. At one locality
Neostethus and Ceratostethus were collected together in large
numbers but neither were collected in association with Phenaco-
stethus. I did not find Neostethus or Ceratostethus in association
with primary freshwater fishes except on one occasion (afternoon,
1 1 July 1970) when Ceratostethus was found in a khlong (strongly
influenced by tides) about midway between Bangkok and Samutsa-
korn, in association with Toxotes, Dermogenys, Rasbora and Eso-
mus. The water was more or less fresh to the taste. A number
of dead Esomus were carried by the current of the khlong, and
a Rasbora or two may have been amongst them. I thought at
the time that mortality of these minnows was perhaps due to in-
cursion of salt water. The dead fish were carried by the out-
going tide.
My impression is that, in Thailand, phallostethids occur in
water that is turbid and fresh; and neostethids in water that is
turbid and brackish or even quite salty. I would guess that at
some of the seven locahties where I collected neostethids the water
was at least a third and perhaps one-half or more as salty as
sea water.
The only phallostethoid in which mating and egg-laying have
been observed (but not in sequence) is the neostethid GulaphaUus
mh-abilis Herre (VilladoUd and Manacop, 1934). It seems prob-
able that all phallostethoids are oviparous. No females carrying
embryos have been observed. It is unclear how much time elapses
between copulation and egg-laying in G. mirabilis, or whether
several egg-layings follow a single copulation. Judging from the
range in size of young individuals found with adults in my collec-
tions of phallostethoids, I suspect that in Thailand species some
reproduction goes on throughout the year. It seems Hkely that
reproductive peaks occur towards the latter part of the rainy sea-
son (November and December).
The distribution of phallostethoids is marginal to the rich East
Indian marine and freshwater fish faunas. They do not occur in
the sea, nor do they penetrate very far into fresh water where
there is a continental fauna of primary freshwater fishes. The
Phallostethidae penetrate further into waters inhabited by primary
freshwater fishes than any other phallostethoids, but they do not
20 BREVIORA No. 374
get very far inland. Usually the canals or creeks they inhabit are
strongly influenced by tides, and thus the faunal composition (with
regard to fishes at any rate) is subject to considerable temporal
variation, involving retreat of primary freshwater fishes and inva-
sion of brackish-water forms, and vice versa. In places where the
tidal changes are greatest, these invasions and counterinvasions
would be a daily event, varying of course with the extent in the
tides. In other places they might occur only during the strongest
tides. Gulaphallus on the island of Luzon probably pass their en-
tire lives in fresh water — where no primary freshwater fishes occur.
DISCUSSION
Selective advantage of internal fertilization in phallostethoids.
One usually associates copulatory organs in teleostean fishes with
viviparity, as in Poecihidae, Embiotocidae, and Brotulidae. It
is a striking fact, however, that several groups of oviparous teleosts
normally have internal fertilization. Among these are some Cot-
tidae, some Scorpaenidae, probably some Clinidae, and probably
some glandulocaudine characids. According to Nelson (1964)
the evolution of internal fertihzation in the oviparous glandulo-
caudine characids of tropical South America appears to be a
response to well-marked wet and dry seasons. The presumed
advantage of a temjxDral separation in mating and spawning is as
follows: mating occurs during the dry season, when populations
are crowded together in small pools, food is scarce and conditions
for survival are generally unfavorable; spawning, on the other
hand, occurs during the height of the rainy season, when the popu-
lation is maximally dispersed (so much so that males and females
may no longer be together) and conditions for survival of the
young are optimum (abundance of food, well-oxygenated waters,
access to areas where most predators are too large to enter).
A very similar set of conditions may apply to the evolution of
internal fertihzation combmed with oviparity in phallostethoids,
with temporal separation of mating and spawning corresponding,
respectively, with periods of low water and high water. This
hypothesis should be extended to the oviparous atheriniform fishes
Horaichthys setnai Kulkarni (1940) and Tomeurus gracilis, males
of which have independently evolved excessively complicated
copulatory organs through modification of the anal fin. Tomeurus
occurs in brackish water along the northeastern coast of South
1971 PHALLOSTETHIDAE 21
America, including the mouth of the Amazon River, Horaichthys
in brackish water along the Bombay and Kerala coast of India.
The sperm in Tomeurus are transmitted in spermozeugmata (Niel-
sen, et al., 1968: 253), as in the viviparous poecilioids (the nature
of the phyletic relationship between Tomeurus and poecilioids is
unclear; they probably are closely related). Horaichthys, on the
other hand, is among the very few teleosts known to have a true
spermatophore (Kulkarni, 1940; Nielsen, et al., 1968). (In true
spermatophores the sperm are enclosed in a capsule. A spermo-
zeugma is an unencapsulated group of sperm held together by
a mucoid substance.)
Trends in the reproductive biology of atherinijorm fishes that
might be conducive to the evohition of internal fertilization. When
oviparous fishes with internal fertilization exhibit a marked delay
between mating and spawning, it may be presumed that fertiliza-
tion actually occurs just before spawning, the sperm having been
stored. Storage of sperm and delayed fertiUzation evidently char-
acterize at least some viviparous fishes, such as those poecihoids
(including forms with superfoetation and forms without it) in
which females are capable of producing up to four or five suc-
cessive broods after a single mating session. It should be noted,
however, that the eggs of a number of atheriniform fishes with
external fertilization normally exhibit arrested embryonic develop-
ment or delayed hatching. A lengthening of the period between
fertilization and hatching, particularly if it could be subjected to
hormonal control, might be highly preadaptive to the evolution
of internal fertilization with delayed spawning. While there is no
evidence of arrested embryonic development or delayed hatching
in atheriniforms with internal fertilization, it seems worthwhile
to review briefly what is known about these phenomena in forms
with external fertilization.
Wourms (1967) found that the eggs of annual cyprinodont
fishes of the subfamily Rivulinae (Cynolebias, Pterolebias, Racho-
via, Nothobranchius, and Austrofundulus) are subject to develop-
mental arrest at one or more stages. In Austrofundulus myersi
dispersion of amoeboid blastomeres occurs early in development
(stages 19-22) so that there is no embryo or aggregation of cells.
This is followed by a slow reaggregation of cells and resumption
of development. This arrested phase (Diapause I) is facultative,
and also occurs in other annuals. Obligate developmental arrests
22 BREVIORA No. 374
in annuals were found in presomite embryos (stage 33, Diapause
II) and in prehatching embryos (stage 44, Diapause III). The
duration of these phases is rather variable. Wourms suggested that
"the net effect of a developmental system which can undergo dia-
pause of variable duration at several stages is to generate a wide
distribution of eggs in any single developmental stage, and to make
the transition from stage to stage a variable phenomenon. The
continued survival of the population is ensured in spite of climatic
cycles which are variable in periodicity and intensity" (Wourms,
1967: 341 1 ). The eggs of annual cyprinodonts have exceptionally
hard chorionic membranes. Harrington (1959) reported delayed
hatching in stranded eggs of marsh killifish, Fundiilus confiuentus,
a member of the cyprinodontid subfamily Fundulinae, and Jones
(1944) reported delayed hatching in Oryzias melastigma, a mem-
ber of the cyprinodontoid family Oryziatidae (see Rosen, 1964,
for a characterization of this family). In O. melastigma hatching
normally occurs in 8-14 days, yet can be delayed up to six weeks;
hatching can be induced by adding water of lower salinity. The
only noncyprinodontoid fishes in which delayed hatching has been
reported as a normal phenomenon are one or two members of the
atherinid subfamily Atherinopsinae, including the grunion, Leu-
resthes tenuis. In this species, "spawning begins just after turning
of tide during the lower (bright moon) series of high tides, but
somewhat later than turn of tide during the higher (dark moon)
series of high tides. Thus the eggs are always deposited near the
same point on the beach profile so that after 2 weeks the lower
series of high tides washes out the eggs deposited during the
higher series, and vice versa. During the interim of low tides, the
eggs are above tide level in moist sand 4 inches below the surface"
(Harrington, 1959: 434-435, after Thompson and Thompson,
1919; Clark, 1925). Grunion eggs presumably have a highly
protective covering. It seems likely that more atheriniform fishes
will be found with either delayed embryonic development or else
deferred hatching (with viability of embryos sustained far beyond
the usual incubation periods). The outer covering of the egg in
many atheriniform fishes is highly protective. These reproductive
features obviously could be advantageous to fishes such as
cyprinodontoids, which generally exploit niches in ephemeral
waters (Foster, 1967).
Finally, it should be noted that Laale and McCallion (1968)
experimentally induced developmental arrest in Brachydanio rerio,
1971 PHALLOSTETHIDAE 23
a cyprinid. Exposure to extracts from Brachydanio, frog, or chick
embryos caused Brachydanio embryos at stages 17-18 to stop
developing. Returned to buffered water, the embryos resumed
normal development and hatched. Laale and McCallion suggested
that the inhibitory factor might be a nuclease.
Foster (1967) suggested that in cyprinodontoids the shift from
expelling and fertilizing all of the ovulated eggs in one continuous
spawning act to expelling and fertilizing them singly or in small
batches greatly increased the amount of courtship behavior per
fertilized egg and thereby the impact of sexual selection. The
habit of expelling a few eggs at a time is also found among ather-
inoids (see Breder and Rosen, 1966) and is apparently char-
acteristic of phallostethoids (Villadolid and Manacop, 1934, for
Gulaphallus mirabilis; Myers, 1935, for Plectrostethus palawanen-
sis) , Horaichthys (Kulkarni, 1940), and Tomeurus (Myers, 1947:
8-11; Breder and Rosen, 1966: 341-343). Whereas in many
atheriniforms the tendency to deposit a few eggs at a time seems
to have led to the evolution of prolonged courtship, in other lines
it may have led to internal fertilization. Supposing females carry
a fair number of eggs but lay only a few at a time, males that can
impregnate the female and fertilize all of the eggs at once have
an obvious selective advantage over males that can fertilize only
a few eggs at a time. Even more to the point, there may be little
to prevent the sperm from getting into the oviduct and fertilizing
eggs retained inside the female. Thus, while fertilization normally
is external in Oryzias, instances of O. latipes females with inter-
nally fertilized eggs (developing embryos) do occur (Amemiya and
Murayama, 1931). Oryzias evidently is ancestral to Horaichthys.
24
BREVIORA
No. 374
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1971 PHALLOSTETHIDAE 25
LITERATURE CITED
Amemiya. I., AND S. MuRAYAMA. 1931. Some remarks on the existence of
developing embryos in the body of an oviparous cyprinodont, Oryzias
{Aplocheiliis) latipes (Temminck et Schlegel). Proc. Imp. Acad. Japan
(Tokyo), 7(4): 176-178.
AuRiCH, H. 1937. Die Phallostethiden (Unterordnung Phallostethoidea
Myers). Intnatl. Rev. Ges. Hydrobiol. Hydrogr., 34: 263-286.
Bailey, R. J. 1936. The osteology and relationships of the phallostethoid
fishes. J. Morph., 59(3): 453-483, 4 pis.
Berg, L. S. 1940. Classification of fishes, both recent and fossil. Trav.
Inst. Zool. Acad. Sci. URSS, 5: 87-517.
Breder, C. M., and D. E. Rosen, 1966. Modes of Reproduction in Fishes.
Garden City, New York, Nat. Hist. Press, xv + 941 pp.
Duncker, G. 1904. Die Fische der Malayischen Halbinsel. Mitt. Naturh.
Mus. Hamburg, 21: 135-207.
Foster, N. R. 1967. Trends in the evolution of reproductive behavior
in killifishes. Stud. Trop. Oceanogr. (Miami), 5: 549-566.
Fowler, H. W. 1937. Zoological results of the third de Schauensee
Siamese Expedition. Part VIII, — Fishes obtained in 1936. Prcc.
Philadelphia Acad. Nat. Sci., 89: 125-308.
Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers.
1966. Phyletic studies of teleostean fishes, with a provisional classifi-
cation of living forms. Bull. Amer. Mus. Nat. Hist., 131, art. 4:
341-455.
Harrington, R. W., Jr. 1959. Delayed hatching in stranded eggs of
marsh killifish, Fimdulus confliientiis. Ecology, 40(3): 430-437.
Herre, a. W. 1925. Two strange new fishes from Luzon. Philippine J.
Sci., 27: 507-513, 2 pis.
1926. Four new Philippine fishes. Philippine J. Sci. 31(4):
533-543, 3 pis.
1939. The senera of Phallostethidae. Proc. Biol. Soc. Wash-
ington, 52: 139-144.
1940. Manacopus, a new name for a genus of Phallosteth-
idae. Copeia, 1940(2): 141.
-. 1942. New and little known phallostethids, with keys to the
genera and Philippine species. Stanford Ichth. Bull., 2(5): 137-156.
HuBBS, C. L., AND L. C. HuBBS. 1945. Bilateral asymmetry and bilateral
variation in fishes. Papers Michigan Acad. Sci. Arts Letters, 30:
229-310, 1 pi.
26 BREVIORA No. 374
Jones, S. 1944. On the occurrence of diapause in the eggs of Indian
cyprinodonts. Current Science (Bangalore), 13: 107-108.
KuLKARNi, C. V. 1940. On the systematic position, structural modifica-
tions, bionomics and development of a remarkable new family of
cyprinodont fishes from the province of Bombay. Rec. Ind. Mus. Cal-
cutta, 42: 379-423.
Laale, H. W., and D. J. McCallion. 1968. Reversible developmental
arrest in the embryo of the zebra-fish, Brachydanio rerio. J. Exp. Zool.,
167(1): 117-127, 3 pis.
Myers, G. S. 1928. The systematic position of the phallostethid fishes,
with diagnosis of a new genus from Siam. Amer. Mus. Novitates, No.
295, 12 pp.
1935. A new phallostethid fish from Palawan. Proc. Biol.
Soc. Washington, 48: 5 6.
1937. Notes on phallostethid fishes. Proc. U. S. Nat. Mus.,
84(3007): 137-143.
1947. The Amazon and its fishes . . . Part 3. Amazonian
aquarium fishes. Aquar. Journ., 18(5) (May, 1947): 6-13, 32.
Nelson, K. 1964. Behavior and morphology in the glandulocaudine fishes
(Ostariophysi, Characidae). Univ. California Pub. Zool., 75(2):
59-152.
NiELSON, J. G., A. Jespersen, and O. Munk. 1968. Spermatophores in
Ophidioidea (Pisces, Percomorphi). Galathea Report, vol. 9: 239-253.
Regan, C. T. 1913. Phallostetlius dunckcri, a remarkable new cyprino-
dont fish from Johore. Ann. Mag. Nat. Hist., 12: 548-555.
1916. The morphology of the cyprinodont fishes of the
subfamily Phallostethinae, with descriptions of a new genus and two
new species. Proc. London Zool. Soc, 1916: 1-26, pis. 1-4.
Rosen, D. E. 1964. The relationships and taxonomic position of the
halfbeaks, killifishes, silversides and their relatives. Bull. Amer. Mus.
Nat. Hist., 127, art. 5: 217-267, pis. 14 and 15.
Smith, H. M. 1927. The fish Neostethus in Siam. Science (n.s.), 65:
353-355.
1945. The fresh-water fishes of Siam, or Thailand. Bull.
U. S. Nat. Mus., No. 188, 622 pp.
TeWinkel, L. E. 1939. The internal anatomy of two phallostethid fishes.
Biol. Bull. Woods Hole, 76(1): 59-69.
ViLLADOLiD, D. v., AND P. R. Manacop. 1934. The Philippine Phal-
lostethidae, a description of a new species, and a report on the biology
of Gulaphallns mirabilis Herre. Philippine J. Sci., 55(3): 193-220,
5 pis.
1971 PHALLOSTETHIDAE 27
WoLTERECK, R. 1942a. Stufen der Ontogenese und der Evolution von
Kopulationsorganen bei Neostethiden (Percesoces, Teleostei). Intnatl.
Rev. Ges. Hydrobiol. Hydrogr., 42: 253-268.
1942b. Neue Organe, durch postembryonale Umkonstruk-
tion aus Fischflossen entstehend. Intnatl. Rev. Ges. Hydrobiol.
Hydrogr., 42: 317-355.
WouRMS, J. D. 1967. A naturally occurring vertebrate dispersion-
reaggregation system subject to developmental arrest. Dissertation
Abstracts, Sect. B, 27(10-11): 3410-3411.
BREVIORA
Miaseiiam of Comparative Zoology
Cambridge, Mass. 15 June, 1971 Number 375
STRUCTURAL HABITATS OF WEST INDIAN ANOLIS
LIZARDS II. PUERTO RICAN UPLANDS
Thomas W. and Amy Schoener^
Abstract. This paper reports differences in structural and climatic
iiabitat between the commonest Anolis species of two upland localities —
Maricao and El Verde — • in Puerto Rico.
Five of the seven Anolis species recorded at Maricao were studied in
detail: two- — cristatelhis and stratitlus — are major components of the
warmer lowland fauna, and three — krugi, evcrmanni, and gundlachi —
are representative of the cooler upland fauna. A. stratiihis and evermanni
tended to occupy higher and thicker perches than the other species; krugi
was found on especially thin perches and often on leaves. The trunk-
crown species evermanni and stratiilus, of very different size, were virtually
totally syntopic over the study area, while the trunk-ground species
cristatelhis and gundlachi, of very similar size, were virtually allopatric.
Similarities and differences in climatic habitat between the species reflected
this microgeography: stratulus and evermanni were found in fairly similar
situations, though the former tended to be in opener areas, while cristatellus
and evermanni were confined to open and closed habitats, respectively. At
El Verde the same species were studied, but the open-area species cristatel-
his and stratulus were relatively rarer. Structural and climatic habitats
paralleled those at Maricao, and the trunk-crown species were again more
syntopic than the trunk-ground species.
In a separate analysis, it is shown that the two trunk-crown species had
significantly more collecting localities in common than did the two trunk-
ground species. The grass-bush species pulchellus and krugi were inter-
mediate in this respect. It is hypothesized that the greater body size
difference of the often syntopic trunk-crown species may be an adaptation
preventing food overlap, an adaptation which the nonsyntopic trunk-ground
species neither have nor need.
1 Biological Laboratories and Museum of Comparative Zoology, Harvard
University, Cambridge, Mass. 02138.
2 BREVIORA No. 375
The possible evolution of climatic and size differences is discussed for
the upland and lowland series. It is suggested that in the broken, drier
forest often found in the lowlands, the trunk-crown species would occur in
more shaded conditions on the average than would the trunk-ground species;
whereas in the wetter, more continuous forests of middle and upper eleva-
tions, the reverse situation would be true. Therefore, the trunk-crown
species may have been predisposed for greater spatial overlap in advance of
their coming together. That the lowland trunk-crown species should be
smaller and less sexually dimorphic in size may be a result of greater aver-
age overlap between it and the corresponding trunk-ground species: possibly
the lower average vegetation height and the greater need to avoid desicca-
tion makes separation by height more difficult in the opener vegetation of
the lowlands. No evidence to support this latter speculation is found in
this study, however.
This is the second in a series of papers describing the structural
habitats of some West Indian Anolis lizards. Rand (1964) has
documented extensively differences between seven Puerto Rican
species in their structural and chmatic habitats. Our study will
also elucidate gross interspecific differences, but its primary pur-
pose is to demonstrate how the size classes of each species studied
are deployed over the vegetation, with respect both to others of
the same species and to similarly sized lizards of other species.
Its secondary purpose is to investigate the spatial relationship be-
tween stratuliis and evermanni, two "trunk-crown" forms (after
Rand and Williams, 1969; Williams and Rand, in preparation)
of very different size and sexual dimorphism, occurring in differ-
ent but overlapping climatic and geographic areas.
FORMAT AND METHODS
Data on the structural habitat were gathered in the way described
previously (Schoener and Schoener, 1971): for each lizard seen
in the area, the type of perch, perch height, and perch diameter
were noted. In addition, when the sun was shining the lizard was
scored as to whether it perched mostly in the sun or shade. Obser-
vations during cloudy weather were lumped into a third chmatic
category. As before, the two of us simultaneously canvassed an
area in staggered fashion so as to minimize the possibility of miss-
ing lizards. In addition, we were assisted during one afternoon at
El Verde by E. E. Williams and W. P. Hall, III.
1971 PUERTO RICAN ANOLIS 3
Results are presented here in two ways: 1) standard, structural
habitat tables are given for each area and lizard class of interest
(Tables 3-7), and 2) the occurrence in sun, shade or clouds is
given for each of the same classes (Table 2).
In order to test for statistically significant differences in struc-
tural and climatic habitat between the hzard classes, the multiway
contingency approach used previously (Schoener, 1970; Schoener
and Schoener, 1971) was again employed. This procedure deter-
mines the strength of associations between the lizard classes and
habitat variables while simultaneously taking into account possible
associations between the habitat variables themselves. It is based
on methods developed by Deming and Stephan (1940), Kullback
(1959), and Bishop (1969), and is exposited for ecologists by
Fienberg (1970); the appendix (page 18) outlines its particular
application to our data. In addition, the appendix contains Tables
9-11, indicating the statistical significance of the comparisons we
make below.
Our presentation begins with a description of the study sites
and their anohne inhabitants, continues with a verbal summary of
differences in habitat between the lizard classes, and terminates in
a comparison of our results to previous ones and in a speculation
on the evolution of size in relation to space for certain Puerto
Rican anoles.
THE SPECIES
Ten species of Anolis are known from Puerto Rico (Table 1).
These can be divided roughly into three groups on the basis of
their climatic and correlated geographic ranges (Rand, 1964; Wil-
liams, Rivero and Thomas, 1965; Heatwole et al., 1969; Web-
ster, 1969).
One group comprises those species which occur modally in rela-
tively humid, dark vegetation at high, cool elevations. This group
consists of 1 ) evermanni — a medium-large, green, trunk-crown
species, relatively sexually dimorphic in size; 2) gundlachi — a
medium-large, rust-brown, relatively dimorphic trunk-ground
species; 3) krugi — a small, striped yellow and brown, relatively
dimorphic grass-bush species; and 4) occultus — a very small,
grey-white, relatively nondimorphic twig species. Two of the
three species, evermanni and krugi, are also found at scattered wet
4 BREVIORA No. 375
lowland localities, probably in remnants of the nearly destroyed
wet lowland forest.
The second group occurs modally in more illuminated, drier
forest at the warmer low to moderate elevations. This group
consists of 1 ) stratulus — a small, grey-brown, relatively non-
dimorphic trunk-crown species; 2) cristatelhis — a medium-large,
brown, relatively dimorphic trunk-ground species; and 3) pulchel-
lus — a small, striped yellow and brown, relatively dimorphic
grass-bush species. This group is also widespread in the highly
disturbed wet lowlands and overlaps the first most commonly at
mesic to wet intermediate altitudes, such as the Maricao locality
to be described below. Additionally, George Drewry (personal
communication) has found cristatelhis predominating in high-
altitude, mossy forest, most of which is above the altitudinal range
of gundlachi.
The third group is restricted to the arid southwestern lowlands
in xeric vegetation. It consists of 1 ) cooki — a medium-large,
grey-brown, relatively dimorphic trunk-ground species, occurring
sympatrically with cristatelhis over much of its range; and 2)
ponceiisis — a small, striped yellow and brown, grass-bush species
of relatively moderate sexual dimorphism. Notice that in this
group there is no trunk-crown representative.
A final species, the nondimorphic green giant ciivieri, appears
from museum records to be commonest at middle and upland ele-
vations, with pockets of abundance in the wetter lowlands such as
those along the north coast.
LOCALITIES
The study to be reported below concentrates on the three
commonest mostly upland species — evennanni, gundlachi and
knigi — and two of the three commonest widespread species —
cristatelhis and stratulus. All five of these species were found in
close proximity to one another in the Maricao Insular Forest, a
preserve in the ''monadnock" region (Pico, 1950) of the Cordillera
Central. Consequently, several adjacent study sites of varying
shapes were marked ofl" in the preserve (altitude = 860-900 m,
slightly less than 4 km due south of the town of Maricao), and
the anoles were observed therein. We distinguished four such areas:
1. Forest interior. This area was the most natural of the four
studied. It contained forest of medium height and somewhat
1971 PUERTO RICAN ANOLIS 5
broken canopy, clinging to a 20-40° mountainside. The under-
story in many places was quite sparse and easily traversed, but
where the canopy had broken, tangled grasses and ferns obstructed
passage. The commonest species seen in this area was gundlachi
(Table 2). The two trunk-crown species, evermanni and stratuliis,
were also common and appeared to be of nearly equal abundance.
In addition, krugi, cristatelliis, and cuvieri occurred as "trace"
species. We saw no occultus though they have been collected there
(Williams, Rivero, and Thomas, 1965).
2. Forest with cleared understory. This area was adjacent to
the more undisturbed forest and consisted of a strip about 30-40
feet wide that bounded an open, grassy parking area. Although
the understory had apparently been cleared, nearly all the large
trees were left standing and very little sun penetrated to the ground,
which itself consisted mostly of bare mud. In this area, gundlachi
was again the most abundantly seen species, followed by evermanni
and stratuliis, again in nearly equal proportions. A few cristatellus
were also seen at the edge of the area.
3. Open forest. This area was less elongated than Area (2)
and faced it from directly across the parking area. Trees were
arrayed in an open, parklike fashion; some overlaid a 2-to-3-foot
herbaceous understory while others grew among short grass. Much
sun penetrated to the ground, in contrast to Area (2). The most
abundant species seen here was cristatellus. Again, evermanni
and stratulus occurred in strikingly similar proportions. A few
gundlachi were seen along the border between this area and Area
(1). In addition, a few krugi inhabited the understory where it
had not been chopped away.
4. Secondary road edge. This area comprised a strip about
1 0-20 feet wide bordering Area ( 1 ) . The understory graded
from a lawnlike growth to dense, tangled typical forest-margin
vegetation. The area faced full sunlight during the morning hours,
that time of day when the sun was most likely to be shining during
the period of our study. Above the understory rose trees of varying
heights, with 40- to 60-foot Cecropia emerging above the rest.
This area showed the most equitable distribution of apparent abun-
dances for the five species. A. stratulus was commonest, but gund-
lachi and evermanni were also rather frequently observed. Less
abundant but far from rare were cristatellus and krugi.
Wc studied the Maricao area during the period June 23-26,
6 BREVIORA No. 375
1969. Weather throughout this time was mostly sunny during the
morning, generally partly cloudy during midday, and somewhat
rainy during the afternoon — on one occasion in the form of a
severe thunderstorm. Rainfall at the nearby but lower town of
Maricao averages 111 inches per year (Pico, 1950).
To further elucidate the relationships between the upland species,
a second, more humid area was investigated at El Verde in the
Luquillo rainforest (340-440 m). The study sites were in parts
of the forest that were and are still the subjects of intensive and
extensive ecological investigation, including a study of its two
commonest Anolis species by Turner, Gist, and Rowland (Odum,
1965). The area is described in detail in various reports (Odum,
1965; Kline, Jordan, and Drewry, 1967, 1968). In comparison
to the Maricao forest, that at El Verde was considerably taller
and of more continuous but still somewhat broken canopy; the
understory was consequently sparser and there were frequent
boulders strewn about the forest floor. Two study sites are dis-
tinguished in Table 2.
The first begins near the biological station and penetrates the
interior of the forest along a trail that climbs upward, eventually
terminating at an observation tower. Our records were all taken
adjacent to the bottom third of the trail where the grade was
closest to horizontal. This area was in coffee plantation until the
early 1930's and had not reached climax but was still in middle
succession at the time of our study (G. E. Drewry, personal com-
munication). The second study site contained roadside secondary
vegetation along the forest margin, very close to the first study area.
In both study areas, gimdlachi was the species most frequently
seen, although it appeared more dominant in the forest interior.
The next most frequently encountered species was evermanni, rela-
tively more abundant along the forest edge. Three other species,
stratulus, knigi, and ciivieh, were rarely encountered in the forest
interior; the first two were always observed in relatively open,
sunny areas. The margin had, however, a more equitable distribu-
tion of species: stratulus, cristatellus and krugi did not appear
uncommon. One cuvieri was also seen. All the relative abundances
given in Table 2 are, of course, apparent only — they represent
what we saw and are therefore biased in favor of three species
{gundlachi, cristatellus, and krugi) frequently occurring at or
below eye level.
Observations were made June 29 -July 1, 1969. Weather dur-
ing this time was, in general, considerably overcast and rainy.
1971 PUERTO RICAN ANOLIS 7
the rain occurring intermittently at all times of the day. Even
though we tried to avoid sunless periods, a heavy incidence of
clouds is apparent in Table 2. Rainfall in the vicinity was esti-
mated from the weekly tables in Kline, Jordan, and Drewry ( 1967,
1968) as 189 inches per year from September 1964 through
August 1966.
RESULTS
Differences between lizard classes. In the following discussion,
all comparisons, unless otherwise stated, are statistically significant
in the sense explained in the appendix. Each species dealt with is
divided into two classes: 1) small individuals, comprising adult
females, subadults, and juveniles, and 2) adult males. The great
majority of the former class (85-100%) were too large to be
labelled "juveniles."
Maricao open areas (Tables 3, 4, 9). In order to compare spe-
cies classes from relatively open areas with respect to their struc-
tural and climatic habitats, observations from the "open forest"
and "secondary road edge" were lumped together. Only the three
commonest species of these areas — evermanni, stratulus and cris-
tatellus — were treated statistically.
Male evermanni perched higher than did all other groups and
were significantly higher than all but male stratulus. The latter
class occurred higher than small evermanni or cristatellus. Small-
sized stratulus were higher than small evermanni and all cristatel-
lus. Small evermanni perched higher than either male or small
cristatellus.
Male evermanni occurred on thicker perches than all other
classes. Male stratulus were on thicker perches than all classes but
male evermanni. Small evermanni were on thicker perches than
small stratulus and small cristatellus. Male cristatellus occupied
thicker perches than did small stratulus.
Within each of the three species, males perched on both higher
and thicker branches than did small-sized individuals.
There were significant differences in the climatic habitats of
stratulus males and the two classes of cristatellus (Table 2). The
former occurred more often in the sun and less often in the shade
than did either of the latter. There was also an overall tendency
for male stratulus to be seen more often on cloudy days. Small
stratulus too occurred more often in the sun than did either class
8 BREVIORA No. 375
of cristatellm and were seen less often in the shade and on cloudy
days. Small-sized evermanni were observed more often in the
shade and less often on cloudy days than were male stratulus;
sunny observations were of about equal proportions in the two
classes. Small evermanni were seen more frequently in the sun
than male cristatellus and less frequently on cloudy days; the per-
cent observations in the shade were nearly identical. Small ever-
manni also occurred more frequently in sunny positions than did
female cristatellus; the latter appeared more often during cloudy
weather and, to a lesser extent, in the shade. No significant climatic
differences were found between the classes of the same species.
Maricao closed area (Tables 5, 6, 10). Observations for two
study sites, the interior forest and the forest with cleared under-
story, were lumped to compare the three most common species —
gundlachi, evermanni and stratulus.
There was no significant difference in perch height between male
stratulus and male evermanni, the two uppermost classes of anoles.
Male evermanni were, however, seen significantly higher than small
stratulus or the classes of gundlachi. Male stratulus did not differ
significantly in height from small evermanni but were observed
higher than either class of gundlachi. Small evermanni were seen
slightly higher than small stratulus and all gundlachi; small stratu-
lus also occurred higher than did the classes of gundlachi.
In perch diameter, male evermanni significantly exceeded all
interspecific classes but male stratulus. Male stratulus, in turn,
significantly exceeded all other interspecific classes but male gund-
lachi. Small evermanni were next in perch diameter, occupying
thicker perches than either class of gundlachi. Small stratulus and
male gundlachi were similar in perch diameter, and the former
occurred on thicker perches than small gundlachi.
Once again, within the same species, the larger classes occurred
on higher and thicker perches than did the smaller-sized classes.
Compared to the other species, both classes of gundlachi
strikingly avoided sunny places. Small gundlachi were found sig-
nificantly more often in the shade or on cloudy days than any
class of stratulus or evermanni. Male gundlachi were found more
often in the shade than small stratulus, and more often in the
shade and on cloudy days than either class of evermanni. As was
the case for the more open area, small evermanni were found
considerably more often in the shade and less often during cloudy
1971 PUERTO RICAN ANOLIS 9
weather than male stratidus; inside the forest, however, they were
also seen slightly more often in the sun. In contrast to the opener
areas, here small evermanni as well differed significantly in the
above respects from small stratidus. No intraspecific comparisons
were significant.
El Verde (Tables 7, 11). Observations for the two areas dis-
tinguished in Table 2 were combined to test structural habitat
differences at El Verde. Only the two most abundant species,
evermanni and gundlachi, were considered.
As before, male and small evermanni were higher than either
class of gundlachi. Male and small evermanni also occupied
thicker perches than did small gundlachi. Male gundlachi, how-
ever, were on greater-diametered perches than small evermanni
and did not differ significantly from male evermanni in this respect,
though the latter at Maricao occupied thicker perches. Intraspe-
cifically, males once again were distributed over greater perch
heights and diameters than small individuals.
There were no significant climatic differences between the lizard
classes — this is unsurprising because of the small sample size
and preponderance of cloudy weather at El Verde.
The common species at El Verde, evermanni and gundlachi,
are similar in size and sexual dimorphism (Table 1) and appear
to stagger their sizes in such a way as to overlap most small ever-
manni and large gundlachi, with respect to both perch height and
perch diameter.
The situation becomes more complex where a third species,
stratulus, is more abundant, as in the Maricao interior forest. The
two species evermanni and gundlachi still maintain the same spatial
relationship between their size classes. However, male stratulus
are now inserted between male evermanni and small-sized ever-
manni. They differ markedly from the former in size (Table 1)
but are only slightly smaller than the latter. There is also rather
strong overlap between small stratulus and small evermanni; the
latter, however, average a good bit larger. Small stratulus also
extensively overlap male gundlachi but are very different in size.
They differ greatly in both perch height and diameter from the
similarly sized small gundlachi.
In the opener areas at Maricao, the situation is essentially the
same for both diameter and height except that cristatellus virtually
replaces gundlachi. Given that substitution, the only difference
10 BREVIORA No. 375
that appears to exist is a reversal in relative height of small stratii-
liis and small evermanni. This reversal allows a more nearly per-
fect staggering of sizes: now male stratulus are further from small-
sized evermanni . All overlaps, of course, are further reduced by
the existence of climatic differences between the niches of the
species.
It is important to point out that in many places in the lowlands
evermanni is absent; there, presumably, cristatelliis and stratulus
often stagger their sizes in much the same way as do evermanni
and gundlachi. However, even if the classes of adult males over-
lapped the most (see below), the two lowland species probably
do not take very similar food since they are so different in size —
in contrast to gundlachi and evermanni. Moreover, it is interesting
to note that where stratulus and evermanni overlap extensively, as
at Maricao, there is only a slightly greater tendency than at "pure"
upland or lowland localities for similarly sized classes to overlap,
as a result of the great difference in size between the two trunk-
crown species.
Because of their small sample size, classes of krugi were not
tested against any other. However, if all observations for krugi
from the three areas are lumped, and to these are added observa-
tions made in high grass and other secondary growth near the mid-
elevation town of Adjuntas, it is possible to compare large and
small krugi in structural habitat. Table 8 shows that males of
krugi occurred significantly higher and on thicker diameters than
did small-sized lizards. However, even more striking was the
much greater frequency of small krugi on leaves. This result is
consistent with those for other species that occupy the leaf habitat
in a major way: carolinensis on Bimini (Schoener, 1968) and
grahami aquarum on Jamaica (Schoener and Schoener, 1971).
No climatic difference was found between the two classes.
Differences between habitat variables (Tables 9-11). In the
Maricao open area the only significant height-diameter associations
were those in which thinner perches tended to be high and thicker
perches tended to be low. Fewer comparisons were significant in
the interior forest at Maricao, but those that were showed a uni-
formly opposite tendency. This could easily reflect our observa-
tional bias: fewer lizards can be seen in the canopy when within
the forest. The one significant interaction at El Verde was the
same as those for the Maricao closed area.
1971 PUERTO RICAN ANOLIS 11
In the open area at Maricao, there was a tendency, when we
combined male-evermanni perches with those of interspecific
classes, for the highest perches to occur in the sun and the lowest
in the shade, as would be expected from the physiognomy of the
vegetation. A somewhat different result was true for the three
significant interactions in the interior vegetation: high perches
were more often used on cloudy days; the lowest perches were
still more frequently found in the shade, however. At El Verde,
the one significant interaction (for male evermanni vs. small gimd-
lachi) was the same as those for the Maricao open area.
There were no significant interactions at all between perch
diameter and insolation in the Maricao closed area, and only one
such interaction was significant for the Maricao open area. This
was a tendency for the combined perches used by small stratulus
and small cristatellus to be thinnest in the sun and thickest on
cloudy days. At El Verde, in contrast, perches of male evermanni
and male giindlachi were significantly thickest in the sun and thin-
nest in the shade. The combined perches for male evermanni and
male gimdlachi also were thinnest in the shade, but the thickest
perches were used during cloudy weather. Both these interactions
may again primarily represent the greater ease of seeing a lizard
on the thin branches and twigs of the understory than on similarly
sized branches belonging to the high canopy of the El Verde rain-
forest. They could, however, also reflect real differences in utili-
zation of the habitat by lizards: for example, during the rather
brief sunny periods at El Verde, lizards may have moved onto
broader surfaces to bask more effectively.
CONCLUSION
Although we have concentrated on differences between the size
classes of the lizards at Maricao and El Verde, our results for
species agree well with those reported by Rand (1964) in his
pioneering study of the structural habitat of species from all areas
of Puerto Rico. In that study Rand pointed out the size difference
between the two trunk-crown species, stratulus and evermanni,
and suggested that, because of their similarity in climatic habitat
(as measured by body and air temperature, as well as shade vs.
sun records), these two species should in general show a greater
amount of spatial overlap than the trunk-ground or grass-bush
12 BREVIORA No. 375
species. Although in an area near the town of Maricao the few
stratulus seen by Rand did not seem to overlap evermanni much
in horizontal habitat, we have shown above that in our areas at
the Maricao Insular Forest, the two species were quite syntopic
and sometimes were of nearly identical apparent abundance (Table
2). Thus we saw 62 evermanni and 66 stratulus in the forest in-
terior, 20 evermanni and 17 stratulus in the forest with cleared
understory, 70 evermanni and 72 stratulus in the open forest, and
36 evermanni and 138 stratulus along the road edge. Even at
our study site at El Verde, where evermanni was seen more fre-
quently (67 evermanni and 7 stratulus in the interior and 31
evermanni and 11 stratulus along the forest margin), stratulus was
interspersed throughout evermanni's horizontal range, although in
the interior it was seen only in fairly open places, such as on
sunny spots along the trail, high on emergent trees, and on stream-
side boulders. G. E. Drewry (personal communication) has, how-
ever, seen stratulus more commonly than evermanni in the more
closed, relatively less leafy canopy of certain other areas at El
Verde.
The extensive syntopy of the trunk-crown species is in striking
contrast to the trunk-ground species we studied. For example, in
the forest interior at Maricao we saw 243 gundlachi and 1 cristatel-
lus. In the open forest at Maricao we saw 175 cristatellus and 8
gundlachi. In the interior forest at El Verde we saw 243 gundlachi
and 0 cristatellus. All three of these areas were relatively equi-
lateral in shape. In the more elongated, marginal areas we saw
more equitable proportions of the two: 33 gundlachi and 6 cris-
tatellus in the forest with cleared understory at Maricao, 50 gund-
lachi and 25 cristatellus along the Maricao secondary road edge,
and 51 gundlachi and 14 cristatellus along the edge locality of El
Verde. At the second of these areas, where the numbers are closest
to identity, we noted that along a strip about ten feet wide cris-
tatellus and gundlachi were perching upon the same vegetation,
although under different climatic conditions: cristatellus was espe-
cially common during sunny days, whereas the majority of the
gundlachi individuals were seen under overcast skies. Thus, even
though there is spatial overlap of cristatellus and gundlachi in
marginal areas, it is largely nonsynchronous.
We can test differences in the relative abundances of trunk-
ground vs. trunk-crown species by constructing 2x2 contingency
1971 PUERTO RICAN ANOLIS 13
tables and running chi-square or exact tests. If this is done for
the six study areas, it is seen that the relative proportions of the
two trunk-ground species are significantly different from those of
the two trunk-crown species in the El Verde interior forest,
Maricao interior forest, Maricao road edge, and Maricao open
forest at the 1 percent level and in the Maricao forest with cleared
understory at the 5 percent level. Only the ecomorphs in the El
Verde forest margin show nonsignificant differences in propor-
tions. In summary, we can conclude that at our study sites the
trunk-ground species were practically allopatric, whereas the
trunk-crown species overlapped broadly.
There exists a second way in which the degree of horizontal
spatial overlap between the species belonging to various habitat
categories may be detected. Museum localities are usually dis-
tinguished horizontally, that is, as points on a map. Therefore, we
have Usted all the localities at which specimens of the six common
Puerto Rican species from the Museum of Comparative Zoology
(Harvard) and the University of Michigan Museum have been
collected. There were 91 such localities for cristateUus and gund-
lachi combined, and only 7 of these (7.7% ) were held in common
between the two species. In contrast, 9 of 45 locahties (20%)
were held in common between evermawu and stratiilus. This pro-
portion was significantly different by a chi-squared test (P < 0.05).
The grass-bush species, kriigi and pulchellus, were intermediate in
this regard: 8 of 59 localities (13.6%) were held in common.
This proportion did not differ significantly from trunk-crown or
trunk-ground species. While the comparisons involve the error
that not all species inhabiting a given locality will be taken by
collectors, usually because of differential abundance and/or acces-
sibility, that error should counteract the pattern brought out in
this analysis. That is to say, because the two trunk-crown species
are seen less often and are more difficult to capture, there is less
likelihood of both being collected at any given locality than both
trunk-ground forms.
It therefore appears clear that the relatively great body-size
difference between the trunk-crown species in Puerto Rico is
associated with their relatively great spatial overlap. Presumably
this size difference in part helps the species to avoid resource com-
petition by being associated with and facilitating differences in the
diet, especially with regard to prey size. However, while the asso-
ciation is presumably adaptive, the mechanism whereby it came
14 BREVIORA No. 375
about is unclear. That is, did the species differ in size to begin
with and so were able to overlap spatially when they came together
to a great degree, or were the size differences largely evolved in
response to a predisposition for spatial overlap in the trunk-crown
forms? We can crystallize the relevant problems by asking three
questions:
1. (a) Why should any of the three structural habitat
groups — trunk-crown, trunk-ground, and grass-bush —
contains species which differ markedly in size? (b) Why
should it be the trunk-crown group, and that only, which
contains such species? There is thus an existence and
uniqueness part to this question.
2. Given that the trunk-crown species show the
greatest size differential, why should the lowland dry-
area form be the smaller, and the upland wet-area form
be the larger?
3. Why should the lowland trunk-crown species be
much less sexually dimorphic in size than the upland
species?
We can gain some insight into the first question by ranking
separately for upland and lowland forest the modal habitats of
the three kinds of species on a darkness or humidity or coolness
gradient (we assume the three to be highly correlated). In middle
and upper elevation forest, such as that at El Verde and Maricao,
the darkest vegetation is the understory, and the most exposed
vegetation lies along margins and in the canopy. Therefore we
would expect the trunk-ground species, which primarily inhabit
the lowest layer of the forest, to be better adapted to dark, cool
conditions than either the trunk-crown species — found largely
in the canopy or somewhat lower — or the grass-bush species,
found mostly along margins. However, the situation is different
for the lowland forest. Much lowland forest in Puerto Rico, par-
ticularly in the south, is and must have been for some time of
xeric aspect. It is characteristic of such forest, as well as of much
coastal woodland elsewhere, to have a relatively broken canopy,
with much light penetrating to the understory and with the largest
trees being often quite dispersed. The patchiness is intensified
during the dry season by some but not all trees undergoing partial
or complete leaf drop. The darkest places for an arboreal lizard
in such veeetation are often associated with the largest trees,
1971 PUERTO RICAN ANOLIS 15
especially often within or directly under the crown. Although that
fraction of the understory immediately beneath such trees will
also usually be shaded, the average degree of shading of the under-
story as a whole should be considerably less than in the humid
forest of middle and upper elevations. The grass-bush habitat
should be even less shaded. Consequently, the trunk-crown habi-
tat should in the lowlands be on the average the darkest and cool-
est; the trunk-ground habitat should be intermediate and the
grass-bush habitat should be on the average the hottest and most
illuminated, even though there are places in these latter two habi-
tats as shaded as any in the crown and on the upper trunk. It
also follows that within the lowland trunk-ground habitat, there
should be a greater diversity of shade regimes than in the upland
trunk-ground habitat. This implies that in the lowlands large
adult males, which prefer thick perches, are more likely to find
such perches shadier than the average small-diametered perch
preferred by the females. This difference should scarcely exist for
the upland species. Therefore, there should be a greater intersexual
difference in climatic habitat for the lowland trunk-ground form
than for the upland trunk-ground form.
Matching species with structural habitat and combining the
upland and lowland species into a single ranking, as would be
appropriate for the many intermediate localities on the island, we
should obtain from sunniest to shadiest: piilchelliis > cristatel-
lus > stratulus >= evermanni = krugi > gundlachi. Notice
that the trunk-ground and grass-bush species are widely separated,
but that the trunk-crown species are adjacent. That is, the
lowland trunk-crown species should occur in the darkest part
of its habitat and the upland trunk-crown species in the sunniest
part of its habitat. Consequently, there should be a great deal of
spatial overlap in intermediate areas on the basis of climatic habi-
tat alone. This means that much of the size difference could
well have evolved as a necessary adjunct to climatic preferenda
of the trunk-crown species, preferenda which have developed inde-
pendently of one another as adaptations to the prevalent vegeta-
tional structure. No such size difference need have evolved for
the trunk-ground or grass-bush species, farther apart on the
climatical spectrum.
The temperature data of Rand (1964, fig. 4), particularly those
for the air, fit for the most part the inequality stated above. Thus,
16 BREVIORA No. 375
among the primarily lowland species, pulchellus occurs at the
warmest temperatures, cristatellus at cooler temperatures, (with
great variance, perhaps because of intersexual differences) and
stratulus the coolest. Almost identical to stratulus is evermanni,
whereas gundlachi is about the same as evermanni and therefore
warmer than predicted, possibly because the highest evermanni
could not be sampled. Values for gundlachi fall below the bulk
of the cristatellus observations. Air temperatures for krugi are
somewhat higher than those for evermanni but markedly below
the majority of the pulchellus observations.
Rand (1964) also provides information on species within five
locahties as to their degree of occurrence in shade. This again
depicts stratulus and evermanni as differing little, while the other
groups differ a good deal. Also shown is that cristatellus and
stratulus can be fairly similar in their shade occurrences, though
in all cases stratulus occupied the deeper shade. A. gundlachi and
evermanni are, however, very different, the former being, of course,
more often in shady positions. In the two localities where both
krugi and pulchellus were found, they showed practically no over-
lap in their shade occurrences.
The climatic data we have gathered at Maricao, which are of
a different sort than those presented by Rand, also support the
greater similarity of the trunk-crown species in their climatic
habitats than the trunk-ground forms. There is not even any
consistent tendency for one or the other of stratulus or evermanni
to be found more often in the sun among the Maricao localities,
although evermanni, if any, appears commoner there. This may
in part, of course, be an artifact of observation; at El Verde, as
we judge from the few stratulus data, the situation appeared to
be reversed. The two trunk-ground species cannot be compared
in most places because one or the other is nearly absent, but along
the secondary road edge, as mentioned, gundlachi was much more
likely to be seen during cloudy weather. At El Verde, despite
the preponderance of overcast skies, cristatellus were seen several
times more frequently in the sun than gundlachi.
A second aspect of the above scheme that can be checked from
our data is the climatic relation between trunk-ground and trunk-
crown forms. Although evermanni and gundlachi showed striking
differences in the expected direction, stratulus and cristatellus did
not. In fact, stratulus were seen significantly more often in the
1971 PUERTO RICAN ANOLIS 17
sun in the open area at Maricao (see above). Possibly this was
an artifact of observation, but it is more likely that in the shade
cristaielliis perches averaged warmer than did those of stratidus
because of the more exposed nature of the vegetation on which
it perched. But Heatwole (1968; Heatwole et ai, 1969) showed
that individuals of stratidus and cristatellus in "an open park-like
situation" at Rio Piedras did not differ significantly in their air
or substrate temperatures. However, Heatwole et al. (1969)
have found that even under apparently identical mean environ-
mental temperatures, the body temperatures of stratulus average
lower than those of cristatellus. Furthermore, cristatellus shows
greater resistance to high temperatures and does not tolerate low
ones as well as stratulus.
Heatwole et al. conclude both from these physiological prop-
erties and Rand's data that "although the two species have similar
ecologies and distributions, in some cases coexisting under appar-
ently identical conditions, there is a tendency for A. cristatellus to
utilize the warmer (less shaded) habitats than A. stratulus." Their
conclusion agrees well with the scheme we have outlined above.
The second question posed above may be related to the relative
degree of overlap between trunk-ground and trunk-crown forms
in the two areas. If we could show that the lowland species are
more likely to overlap spatially than the upland forms, then we
could argue that the smaller size of stratulus may have evolved in
part as a response to that overlap and thus served as a preadapta-
tion for any further displacement that might have occurred between
the trunk-crown species. Rand's structural habitat grids show
practically no differences in percent overlap (62.5% vs. 61%)
between the two species pairs. However, his data were from sev-
eral localities and all size classes combined. Even if only adult
males are considered and our data for particular locahties used,
results are inconsistent: percent overlap for evermanni and gund-
lachi is 35 percent in the forest interior at Maricao and 72 per-
cent at El Verde, whereas that for stratulus and cristatellus is 30
percent along the Maricao road and 50 percent in the Maricao
open forest. It is possible that these results are artifacts of the
greater difficulty of seeing a trunk-crown species in the canopy
when within a continuous forest than when along a forest margin
or in a very open area. That error would tend to minimize differ-
ences between evermanni and gundlachi and is almost certainly
18 BREVIORA No. 375
in part responsible for the high overlap value at El Verde. One
might argue, a priori, that because of the greater average height
of rainforest such as that at El Verde, overlap between two spe-
cies segregated mainly by height should be less than in drier forest.
Furthermore, even though the species are physiologically adapted
to different thermal environments, it is possible that in the dry
lowlands the greater danger of desiccation forces cristatelliis and
stratuliis together in shaded situations during a large portion of
the day; in the uplands, however, no such problem need arise,
and the more exposed species, in this case evermarmi, can occur
in sunny or open places most of the day. This is perhaps the rea-
son why less difference was found between the climatic habitats
of cristatellus and stratuliis than between gnndlaclii and evermanni
in our study. Any overlap between the first pair should be espe-
cially severe for male cristatellus because they more frequently
occupy the relatively large trees which stratulus inhabit.
If the explanation above be correct, that is, had stratulus evolved
small size in part to alleviate competition with male cristatellus,
then the reduced sexual dimorphism of stratulus would automati-
cally follow, since it is the male that would be displaced.
ACKNOWLEDGMENTS
We thank G. E. Drewry, A. S. Rand, T. P. Webster, and E. E.
Williams for critical comments on the manuscript and S. D. Fien-
berg for statistical advice. We also thank G. E. Drewry for his
hospitality at the Puerto Rican Nuclear Center field station at
El Verde and K. Horton for typing the tables photographed directly
for this article. Rand, Webster and Williams were all in the field
with us during some part of this study, and their previous experi-
ence with the island greatly facilitated our work. In addition,
Williams and W. P. Hall III contributed some observations to our
data from El Verde. Research was partly supported by NFS
grants GB 6944 and B 01 980 IX to E. E. Williams.
STATISTICAL APPENDIX
This appendix briefly reiterates the description of the statistical
treatment of the structural and climatic habitat data given in detail
elsewhere (Schoener and Schoener, 1971). Four variables —
lizard class, climatic category, perch height and perch diameter —
were used to set up 4-way contingency tables. For the lizard and
1971 PUERTO RICAN ANOLIS 19
climatic variables, two (the classes being compared) and three
(sun, shade, and clouds) categories (levels), respectively, were
used. For perch height and diameter, two levels each were chosen
by separately determining the point of maximum difference in
cumulative frequency between the distributions of the habitat
variable for the two lizard classes being compared. Data were
broken at this point, all observations less than or equal to that
number being cast into one category and all greater than that
number being cast into the other. Table 12 lists critical values
for height and diameter groupings. As before, an iterative pro-
cedure (Deming and Stephan, 1940; Bishop, 1969; Fienberg,
1970) was used to fit the data to models containing all six 2-way
interactions. One by one, interactions were dropped if differences
between models were not significant at the 5 percent level accord-
ing to difference in the log-likelihood ratio chi-square (Kullback,
1959; Ku and Kullback, 1968). Results are summarized in
Tables 9-1 1 . The following key applies to the numbers in the
body of the table:
1 — the interaction was significant every time it was tested in
the removal procedure;
2 — the interaction was significant at least at the termination
of the procedure;
3 — the interaction was significant when removed from the most
inclusive model (with six interactions) but not at termination;
4 — the interaction was significant sometime during the pro-
cedure but not at the beginning or end;
0 — the interaction was never significant.
As can be seen, nearly all interactions could be labelled "0" or
"l." There was no set of models for a particular 4-way table
which never gave a x" value denoting a satisfactory fit of the model
at the 5 percent level, regardless of what interactions were re-
moved. Therefore we did not test for 3-way interactions. There
were only three tables with zero margins. These could be, and
therefore were, handled in the way given by Fienberg (1970).
LITERATURE CITED
Bishop, Y. M. M. 1969. Full contingency tables, logits, and split con-
tingency tables. Biometrics, 25: 383-400.
Deming, W. E., and F. F. Stephan. 1940. On a least squares adjustment
of a sampled frequency table when the expected marginal totals are
known. Ann. Math. Stat., 11: 427-444.
20 BREVIORA No. 375
FiENBERG, S. E. 1970. The analysis of multidimensional contingency
tables. Ecology, 51: 419-433.
Heatwole, H. 1968. Relationship of escape behavior and camouflage in
anoline lizards. Copeia, 1968: 109-113.
, T. Lin, E. Villalou, A. Muniz, and A. Matta. 1969.
Some aspects of the thermal ecology of Puerto Rican anoline lizards.
J. Herpetology, 3: 65-78.
Kline, J. R., C. F. Jordan, and G. E. Drewry. 1967, 1968. The rain
forest project annual reports. Puerto Rico Nuclear Center, Puerto Rico.
Ku, H. H., and S. Kullback. 1968. Interactions in multi-dimensional
contingency tables: an information theoretic approach. J. Res. Natl.
Bur. Standards - Mathematical Sciences, 728: 159-199.
Kullback, S. 1959. Information Theory and Statistics. Dover Publica-
tions, New York.
Odum, H. T. 1965. The rain forest project annual report FY-65. Puerto
Rico Nuclear Center, Puerto Rico.
Pico, R. 1950. The geographic regions of Puerto Rico. Univ. Puerto Rico
Press. Rio Piedras, Puerto Rico. 256 pp.
Rand, A. S. 1964. Ecological distribution in anoline lizards of Puerto
Rico. Ecology, 45: 745-752.
, and E. E. Williams, 1969. Anoles of La Palma: aspects of
their ecological relationships. Breviora, No. 327: 1-19.
Schoener, T. W. 1968. The Anolis lizards of Bimini: resource partition-
ing in a complex fauna. Ecology, 49: 704-726.
. 1969. Size patterns in West Indian Anolis lizards. I. Size
and species diversity. Syst. Zool., 18: 386-401.
1970. Nonsynchronous spatial overlap of lizards in
patchy habitats. Ecology, 51: 408-418.
, AND A. Schoener. 1971. Structural habitats of West
Indian Anolis lizards. I. Lowland Jamaica. Breviora, No. 368: 1-53.
Webster, T. P. 1969. Ecological observations on Anolis occult us Wil-
liams and Rivero (Sauria, Iguanidae). Breviora, No. 312: 1-5.
Williams. E. E.. and A. S. Rand. The structure of diversity in Anolis liz-
ards, in prep.
, J. A. RivERO, AND R. Thomas. 1965. A new anole
(Sauria, Iguanidae) from Puerto Rico. Breviora, No. 231: 1-18.
1971 PUERTO RICAN ANOLIS 21
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to
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97
PUF.RTO RICAN ANOI.IS
23
Table 2. cliiratic and geographic occurrence of Puerto Rican anoles.
Percent Occurrence
Lizard Class
Sun
Shade Clouds
Maricao, forest interior
evermanni
adult
male
everraanni
small
gundlachi
adult
male
gundlachi
small
stratulus
adult
male
stratulus
small
cristatellus
adult
male
crista tellus
small
krugi
adult
male
krugi
small
cuvieri
large
aricao, forest, clea
evermanni
adult
male
evermanni
small
gundlachi
adult
male
gundlachi
small
stratulus
adult
male
stratulus
small
cristatellus
adult
male
cristatellus
small
krugi
adult
male
I-.rugi
small
cuvieri
large
53.8
11.5
50.0
28.6
0
0
33.3
9.1
66.7
16.7
57.1
25.0
16.0
0
18.2
33.3
Sun or Sample
Shade Size
34.6
44
.4
22
.2
27.8
17
1
34
.3
45.7
15
6
32
4
48.0
31
8
4
5
63.6
31
8
13.
6
47.7
0
0
100.0
50.0
83.3
14.3
75.0
84.0
66.7
72.7
5.6
2.9
4.0
0
6.8
0
26
36
70
173
22
4 4
1
0
2
0
1
14
8
25
6
11
0
6
0
0
0
Percent
Compo-
sition
6.9
9.6
18.7
46.1
5.9
11.7
0.3
0.5
0.3
7.9
18.4
10.5
32.9
7.9
14.5
7.9
24
BREVIORA
No. 375
ass
Percent
Occurrence
Sample
Size
Lizard CI
Sun
Shade
Clouds
Sun or
Shade
Percent
Compo-
sition
Maricao, open
forest
evermanni
adult
male
33.3
28.6
33.3
4.8
21
6.2
evermanni
small
44.9
28.6
20.4
6.1
49
14.5
gundlachi
adult
male
0
gundlachi
small
12.5
50.0
25.0
12.5
8
2.4
stratulus
adult
male
40.5
8.1
45.9
5.4
37
11.0
stratulus
small
22.9
37.1
37.1
2.9
35
10.4
crista tellus
adult
male
22.0
26.0
50.0
2.0
50
14.8
cristatellus
small
22.4
32.0
41.6
4.0
125
37.1
krugi
adult
male
28.6
42.9
28.6
0
7
2.1
krugi
small
0
60.0
40.0
0
5
1.5
cuvieri
large
0
Maricao, secondary road
edge
evermanni
adult
male
78.3
0
21.7
0
23
8.3
evermanni
small
69.2
7.7
23.1
0
13
4.7
gundlachi
adult
male
11.1
27.8
61.1
0
18
6.5
gundlachi
small
15.6
3.1
78.1
3.1
32
11.5
stratulus
adult
male
55.6
7.4
33.3
3.7
54
19.4
stratulus
small
59.5
8.3
31.0
1.2
84
30.2
cristatellus
adult
male
33.3
33.3
16.7
16.7
6
2.2
cristatellus
small
36.8
21.1
31.6
10.5
19
6.8
krugi
adult
male
0
25.0
50.0
25.0
4
1.4
krugi
small
4.0
16.0
76.0
4.0
25
9.0
cuvieri
large
0
El Verde, interior
evermanni
adult
male
5.6
33.3
61.0
0
18
5.6
evermanni
small
3.9
27.6
68.6
0
51
15.7
gundlachi
adult
male
7.1
32.5
59.8
0.6
169
52.2
gundlachi
small
5.4
41.9
52.7
0
74
22.8
1971 PUERTO RICAN ANOLIS 25
stratulus
adult
male
100.0
0
0
0
1
0.3
stratulus
small
33.3
33.3
33.3
0
6
1.9
cristatellus
adult
male
0
cristatellus
small
0
krugi
adult
male
50.0
0
50.0
0
2
0.6
krugi
small
0
0
100.0
0
4
1.2
cuvieri
small
0
0
100.0
0
1
0.3
El Verde, edge
evermanni
adult
male
46.2
23.1
30.8
0
13
9.5
evermanni
small
27.8
16.7
55.6
0
18
13.1
gundlachi
adult
male
8.3
36.1
52.8
2.8
36
26.3
gundlachi
small
13.3
26.7
60.0
0
15
10.9
stratulus
adult
male
67.7
0
33.3
0
3
2.2
stratulus
small
37.5
12.5
37.5
0
8
5.8
cristatellus
adult
male
50.0
0
50.0
0
6
4.4
cristatellus
small
25.0
25.0
50.0
0
8
5.8
krugi
adult
male
16.7
16.7
66.7
0
6
4.4
krugi
small
21.7
26.1
52.2
0
23
16.8
cuvieri
small
100.0
0
0
0
1
0.7
26
BREVIORA
No. 375
Table 3. Maricao secondary road edge. Percent observations in
various structural habitat categories. H = >20'; G = ground; R =
rocks; N = sample size.
^^^^iameter
Ht. ^-^in.)
(feet) ^-v,^^^
>5
5-2 1/2 2
1/4-7/8
7/8-1/8
le
aves
Total
N = 54
male
stratulus H = 11
G = 0 R =
• 0
10.5-20
4
17
4
0
0
25
5-10
9
17
6
0
0
32
3-4 3/4
9
4
4
0
0
17
<3
9
4
4
0
0
17
Total
31
42
18
0
0
N = 84
small
stratulus
H = 2
G = 0 R =
= 0
10.5-20
4
8
7
1
0
20
5-10
4
13
13
6
0
36
3-4 3/4
2
6
5
5
0
18
<3
10
6
6
1
2
23
Total
20
33
31
13
2
N = 6
male
cristatellus
H = 0
G = 0 R
= 0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
0
0
0
3-4 3/4
17
17
0
0
0
34
<3
33
17
17
0
0
77
Total
50
34
17
0
0
N = 19
small
cristatellus H = 0
G = 11
R =
0
10.5-20
0
0
0
0
0
0
5-10
0
0
0
0
0
0
3-4 3/4
5
0
0
0
0
5
<3
16
26
26
16
0
84
Total
21
26
26
16
97:
PUERTO RICAN ANOLIS
27
N = 23
male
everinanni
H
= 4
G = 0
R =
0
10.5-20
22
17
0
0
0
39
5-10
9
13
0
0
4
26
3-4 3/4
17
0
0
0
0
17
<3
13
0
0
0
0
13
Total
61
30
0
0
0
N = 13
small
evermanni
H
= 0
G = 0
R =
0
10.5-20
0
8
0
0
0
8
5-10
0
15
0
0
8
23
3-4 3/4
8
0
0
0
0
8
<3
46
15
0
0
0
61
Total
54
38
0
0
8
Table 4. Maricao open forest. Percent observations in various
structural habitat categories. H = >20'; G = ground; R = rocks; N =
sample size.
JDiameter
Ht. ^\^(in.)
(feet)
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
N = 37
10.5-20
5-10
3-4 3/4
<3
Total
male stratulus
H
= 0
G =
= 0
R =
= 0
0 22
0
0
0
8 16
11
5
0
3 3
0
5
0
5 19
3
0
0
22
40
11
27
14
10
N = 35
10.5-20
5-10
3-4 3/4
<3
Total
small stratulus
H =
0
G =
= 0
R =
= 0
0 3
0
0
0
3 6
19
13
0
9 0
11
9
0
14 9
6
0
0
3
41
29
29
26
18
36
22
28
BREVIORA
No. 375
N = 50
10.5-20
5-10
3-4 3/4
<3
Total
•M = 12 5
10.5-20
5-10
3-4 3/4
<3
Total
N = 21
10.5-20
5-10
3-4 3/4
<3
Total
male
cristatellus
H =
= 0
G
= 6
R =
= 0
0
6
0
0
0
4
4
10
2
0
8
0
12
2
0
.20
10
7
7
2
32
20
29
11
small
crista
te
llus
H =
0
G =
12
R
= 1
0
2
0
0
0
0
2
5
2
0
2
2
2
3
1
11
23
16
15
2
13
29
23
20
male
evermanni
H
= 0
G =
= 10
R =
0
5
19
7
2
0
5
5
14
5
0
10
5
0
0
0
14
0
0
0
0
6
20
22
46
2
9
9
67
33
29
15
14
34
29
21
N = 49
10.5-20
5-10
3-4 3/4
<3
Total
small
evermanni
H =
6
G
= 2
R =
0
2
6
2
4
0
4
6
12
10.
0
0
2
4
0
0
16
10
8
2
2
14
32
6
38
22
24
26
16
1971
PUERTO RICAN ANOLIS
29
Table 5. Maricao forest with cleared understory. Percent
observations in various structural habitat categories. H = >20'
G = ground; R = rocks; N = sample size.
^iameter
Ht. ^^-.^^in.)
(feet)
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
N = 6
10.5-20
5-10
3-4 3/4
<3
Total
male
stratulus
H
= 50
G =
0
R = 0
0
17
0
0
0
17
17
0
0
0
0
0
0
0
0
0
0
0
0
0
17
34
0
0
N = 11
10.5-20
5-10
3-4 3/4
<3
Total
small
stratulus
H
=
0
G =
0
R = 0
0
9
0
0
0
9
18
9
9
0
0
0
0
0
0
18
18
9
0
0
9
45
0
45
27
45
N = 8
10.5-20
5-10
3-4 3/4
<3
Total
male gundlac
hi
H
= 0
G =
= 0 R =
0
0 0
0
0
0
0 12
12
25
0
0 12
0
12
0
0 0
19
6
0
0
49
24
25
24
31
43
N = 25
10.5-20
5-10
3-4 3/4
<3
Total
small gundlachi
H =
0
G =
= 0
R =
= 0
0 0
0
0
0
4 4
4
4
0
0 16
8
4
0
4 12
28
12
0
0
16
28
56
32
40
20
30
BREVIORA
No
. 375
N = 6
male
evermanni
H = 0
G = 0 R = 0
10.5-20
17
0
0
0
17
34
5-10
0
33
17
0
0
50
3-4 3/4
0
0
0
0
0
0
<3
0
17
0
0
0
17
Total
17
50
17
0
17
N = 14
small
evermanni
H = 0
G = 7 R = 0
10.5-20
0
0
0
0
0
0
5-10
7
14
14
0
0
35
3-4 3/4
0
0
0
0
0
0
<3
36
21
0
0
0
57
Total
43
35
14
0
0
Table 6. Maricao forest interior,
various structural habitat categories,
rocks; N = sample size.
Percent observations in
H = >20'; G = ground; R =
^\Diameter
Ht. ^\(in.)
(feet) ^v^
>5
5-2 1/2
2 1/4-7/8 7/8-1/8
leaves
Total
N = 22
male
stratulus
H = 5 G = 0
R =
0
10.5-20
0
32
5 0
0
37
5-10
5
14
7 11
5
42
3-4 3/4
0
0
9 0
0
9
<3
5
5
0 0
0
10
Total
10
51
21 11
5
N = 44
small
stratulus
H = 2 G = 2
R
= 0
10.5-20
2
2
2 2
2
10
5-10
5
5
16 11
7
44
3-4 3/4
0
0
2 2
0
4
<3
2
9
20 2
2
35
Total
16
40
17
11
1971
PUERTO RICAN ANOLIS
31
N
= 70
10.5-20
5-10
3-4 3/4
<3
Total
N
= 173
10.5-20
5-10
3-4 3/4
<3
Total
N
= 26
10.5-20
5-10
3-4 3/4
<3
Total
N = 36
10.5-20
5-10
3-4 3/4
<3
Total
male gundlachi
H
= 0
G =
1 R = 0
0 0
0
0
0
4 4
6
4
0
1 11
13
7
1
9 11
11
13
1
14
26
small gundlachi
0 0
1 1
0 1
3 9
30
H = 0
0
0
5
23
G =
24
8
0
3
4
29
R = 1
0
0
1
11
35
24
20
4 11
28
36
12
male evermanni
H
= 8
G =
= 0 R =
= 0
15 4
8
0
0
12 12
12
8
0
0 8
0
0
8
8 0
0
0
0
0
18
33
45
0
5
11
75
27
44
16
small
evermanni
H =
3
G
= 3
R =
0
0
8
11
0
0
0
6
17
14
3
0
3
6
3
3
0
6
8
8
0
19
40
15
24
23
42
25
32
BREVIORA
No. 375
Table 7. El Verde. Percent observations in various structural
habitat categories. H = >20'; G = ground; R = rocks; N = sample size.
^^"'----.Diaineter
Ht.^^^in.)
(feet) ^^..^^
>5
5-2 1/2
2 1/4-7/0
7/8-1/8
leaves
Total
N = 205
male
gundlachi
H = 0 G = 2
R =
1
10.5-20
0
3
0
0
0
3
5-10
15
10
12
7
0
44
3-4 3/4
8
12
9
3
0
32
<3
9
4
2
1
0
16
Total
32
29
23
11
0
N = 89
small
gundlachi
H = 0 G
= 4
R
= 3
10.5-20
0
0
0
0
0
0
5-10
0
8
8
6
1
23
3-4 3/4
4
5
9
10
0
28
<3
6
12
9
12
0
39
Total
10
25
26
28
1
N = 31
male
evermcinni
H = 0 G =
= 3
R =
0
10.5-20
6
0
6
0
0
12
5-10
16
3
10
6
3
38
3-4 3/4
13
3
13
0
0
29
<3
10
3
3
0
0
16
Total
45
9
32
6
3
N = 69
small
evermanni
H = 1 G
= 6
R
= 3
10.5-20
6
1
0
1
1
9
5-10
1
4
12
9
6
32
3-4 3/4
4
3
3
9
0
19
<3
13
9
2
4
0
28
Total
24
17
17
23
7
1971
PUERTO RICAN ANOLIS
33
Table 8. A. krugi, all localities combined. Percent observa-
tions in various structural habitat categories. H = >20'; G = ground;
R = rocks; N = sample size.
^iameter
Ht. ^\(in.)
(feet)
>5 5-2 1/2 2 1/4-7/8 7/8-1/8 leaves Total
N = 33
10.5-20
5-10
3-4 3/4
<3
Total
N = 83
10.5-20
5-10
3-4 3/4
<3
Total
male krugi
0 3
0 0
0 0
0 0
H = 0 G
0
6
12
3
0
R = 0
3
30
6
12
21
51
0
15
6
3
24
small krugi
H =
0
G =
4
R = 0
0 0
0
0
0
0 0
0
1
5
0 0
0
7
16
0 0
2
11
54
6
51
24
18
0
6
23
67
19
75
34
BREVIORA
No. 375
w
G
O
U)
M
e
o
u
(fl
(1)
to
C
0)
o
0
«3
U
■M
m
s
u
o
O
o
c
(0
o
•H
c
en
■H
n3
o
•H
tn
-p
en
0)
en 0)
>
0)
ITS
tn
0
i-\
rH
0)
rH
+J
iH
fl
(0
+J
e
tn
U)
■H
i-i
u
tn
:3
iH
tH
(U
0)
iH
+J
03
(0
g
-M
tn
•H
M
U
H H H
* * *
O <H r-H iH
rH
CO
en
3
3
rH
CO
r-i
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1971
PUERTO RICAN ANOLIS
35
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36
BREVIORA
No. 375
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1971
PUERTO RICAN ANOLIS
37
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38
BREVIORA
No. 375
Table 11. Statistical significance for El Verde."
^^^^Group vs.
Gro^i^^i^ht
vs. ^""~\^
male
small
male
small
Diameter ^""^
evermanni
evermanni
gundlachi
gundlachi
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1
2
1
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1*
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1
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0
1
1
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1*
1*
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Hei^h^-^°1^^^°'^
vs. ^"^-^^
diameter ^~^--^^^
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0
0
0
small evermanni
0
0
0
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0
0
0
small gundlachi
1(A)
0
0
^""--.^eight vs.
^^^^insolaticn
Diameter^^^^^
vs. ^^-^^^
insolation ^~^^
-^
male evermanni
0
0
1(F)
small evermanni
0
0
0
male gundlachi
1(K)
0
0
small gundlachi
2(L)
0
0
" * = species at top has larger value; A = small diameters at low perches;
F = highest in sun and lowest in shade; K =: thickest in sun and thinnest
in shade; L r= thinnest in shade and thickest in clouds: for interpretation
of numbers, see "Statistical appendix."
1971
PUERTO RICAN ANOLIS
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BREVIORA
Mmseiuinii of Coiniiparative Zoology
Cambridge, Mass. 15 June, 1971 Number 376
Podocnemis venezuelensis, a new fossil pelomedusid (Tes-
tudines, Pleurodira) from the Pliocene of Venezuela and
a review of the history of Podocnemis in South America
Roger Conant Wood^
and
Maria Lourdes Diaz de Gamero-
Abstract. Podocnemis venezuelensis, a new species of pelomedusid
turtle from the mid-Pliocene (Huayquerian) of Venezuela is described.
Present evidence is insufficient for determining whether it was a marine
or a fresh water form. Previously described South American fossil species
that have been attributed to Podocnemis are discussed. Of these, only
two — P. bassleri and P. elegans — are surely referable to this genus. It
is concluded that the fossil record of Podocnemis is not yet adequate for
reconstructing its evolutionary history on this continent.
Resumen. Se describe una nueva especie de tortuga pelomedusida
del Plioceno medio (Huayqueriense) de Venezuela, Podocnemis venezu-
elensis. Las presentes pruebas son insuficientes para determinar si fue de
aguas marinas o de agua dulce. Se discuten las especies suramericanas
fosiles que han sido atribuidas a Podocnemis. De estas, solo dos — P. bass-
leri y P. elegans — son valederas. Se concluye que el registro fosil de
este genero no es aiin adecuado para la reconstrucion de su historia evo-
lutiva en este continente.
INTRODUCTION
Only one species of fossil turtle, Podocnemis geologorum
(Simpson, 1943), has ever been formally described from Vene-
zuela. Other occurrences of Venezuelan fossil chelonians have,
1 Museum of Comparative Zoology, Harvard University.
2 Escuela de Geologia y Minas, Universidad Central de Venezuela,
Caracas.
2 BREVIORA No. 376
however, occasionally been mentioned: Swinton (1928) referred
three specimens "too fragmentary for specific determination," of
Pliocene or Pleistocene age (Van Frank, 1957: 22n), to "Testuclo
sp."; and Royo y Gomez (1960) published a faunal list of fossil
vertebrates discovered in what is now known as the upper member
of the Urumaco Formation, that included ". . . varios capara-
zones completos . . . de tortugas palustres (Podocnemys) . . . ."
which he believed to be of mid-Miocene age. Unfortunately, the
untimely death of Royo y Gomez in 1961 prevented his describing
any of this material. The specimens, however, are preserved in
the collections of the Universidad Central de Venezuela in Ca-
racas. Except for one report on rodent remains (Pascual and Diaz
de Gamero, 1969) and another on a new gavial (Sill, 1970),
nothing has been published on them. The fossil turtles are de-
scribed here.
Abbreviations used in this paper are:
AMNH — the American Museum of Natural History, New York
MCN — Museo de Ciencias Naturales, Caracas
VF — Laboratorio de Paleontologia, Escuela de Geologia de la
Facultad de Ingeniera de la Universidad Central de Venezuela
(Caracas).
CLASSIFICATION AND DESCRIPTION
Order Testudines
Suborder Pleiirodira
Family Peloiiiedusidae
Genus Podocnemis
A cautionary word is appropriate concerning the attribution of
fossil material to Podocnemis. Most extinct species of this genus
are known solely from shell material. However, at least one other
fossil genus, Botliremys, has a shell that is virtually indistinguish-
able from that of Podocnemis. In fact, two purported species of
Podocnemis, P. barberi (Schmidt, 1940) and P. alabamae (Zan-
gerl, 1948), were originally described on the basis of shell material
that, upon the later discovery of associated skull material, has
subsequently been referred to the genus Botliremys (GafTney and
Zangerl, 1968). Conceivably, therefore, when skulls become
known for some of the fossil species now referred to Podocnemis
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 3
on the basis of their shells, it may be necessary to transfer them
elsewhere. The new species described in this paper is placed in
Podocnemis with this caveat.
Podocnemis venezuelensis sp. nov.
(Figures 1-3 and Plates I-V )
Type. VF 1176, a complete carapace, slightly flattened, includ-
ing much of the axial buttresses and the pelves, the latter badly
damaged. Also, VF 1173, a fairly complete but badly fractured
plastron lacking only the entoplastron and epiplastra. Although
bearing a diff'erent number, the plastron is of an appropriate size
and state of preservation so that we believe it highly probable
that both of these shell elements are from the same individual
(see Plate IV). Unfortunately, none of Royo y Gomez's notes
contain information bearing on this point.
Hypodigm. The type, and VF 1174, a plastron lacking the
posterior lobe; VF 1175, the anterior one-third of a carapace; VF
1177, a complete carapace and plastron, considerably flattened,
of a relatively small individual; VF 1177a, a nearly complete
carapace and plastron of the largest individual in the sample,
lacking only the pygal and posterior portions of both eleventh
peripherals; VF 1177b, the left epiplastron, hyoplastron, and left
one-half of the entoplastron, plus several pleurals attached to part
of the bridge, probably all belonging to the same individual; VF
1059, the distal end of a right humerus.
Horizon and locality. The "capa de huesos" or "capa de tor-
tugas" (Pascual and Diaz de Gamero, 1969: 373 and map) of
the upper member of the Urumaco Formation, probably of mid-
Pliocene (Huayquerian) age, north of Campo El Mamon, state
of Falcon, Venezuela.
The Uramaco Formation is divided into three members, desig-
nated as lower, middle, and upper. The lower and middle mem-
bers are largely, if not entirely, of marine origin while the upper
member apparently includes both littoral or deltaic as well as more
strictly terrestrial facies. Marine molluscs from the middle mem-
ber were the basis for the original age determination of mid-
Miocene for this formation. This was subsequently revised to late
Miocene. However, the recent study of Pascual and Diaz de
Gamero (1969: 379) indicates that at least the "bone bed" of
the uppermost part of the upper member is probably of middle
BREVIORA
No. 376
Figure. 1 Reconstruction of the shell of Podocnemis venezuelensis
(dorsal view). The scale represents a length of 15 centimeters. The posi-
tions of the axial and inguinal buttresses as well as of the iliac scars of
the pelvis on the visceral surface of the carapace are indicated by the
broken lines.
1971
NEW VENEZUELAN FOSSIL PELOMEDUSID
Figure 2. Reconstruction of the shell of Podocnemis veneziielensis
(ventral view). Same scale as in Fig. 1. The positions of the pelvic scars
on the visceral surface of the plastron are indicated by the broken lines.
6 BREVIORA No. 376
Pliocene (Huayquerian) age. Palynological evidence appears to
confirm this date (A. E. Gonzalez Guzman, personal communica-
tion). It is possible that there may be a temporal hiatus between
the middle and upper members of the Urumaco Formation that
cannot be detected structurally; an explanation of this nature
would account for the different age determinations of the middle
and upper members of the Urumaco Formation.
Diagnosis. Differs from all other species of Podocnemis in
totally lacking neural bones.
Description. It is unfortunate that a gypsiferous encrustation
on the external surfaces of all of the specimens obscures most of
the bone sutures and virtually all scute sulci. Nevertheless, an
essentially complete osteological description of the shell is possible
from examination of the visceral surfaces of the type and some
other, less complete specimens. Although the outlines of any par-
ticular bone may differ somewhat from the external to the internal
surface of the shell, as Dacque (1912: 290 and fig. 7) has demon-
strated, it is nonetheless possible to make a reasonable reconstruc-
tion of the shell of a typical representative of the species (see
Figures 1 and 2 ) .
Adult specimens of Podocnemis venezuelensis attained a rather
large size. The three complete carapaces in our sample range
from approximately 46 to 67 centimeters' in length. Of the eight
living species of Podocnemis, only two, P. expansa and P. unifilis,
are reported to reach a larger maximum size.
In cross section, the arch of the carapace is very flat; in outline
it is somewhat oval and greatly expanded posteriorly. The external
surface of the carapace is completely smooth. No significant in-
dentation occurs in the nuchal region. Well-developed axial and
inguinal buttresses unite the carapace to the plastron. The plastron
itself is essentially flat and of nearly uniform thickness throughout.
The outstanding feature of the carapace, and indeed the char-
acter permitting definition of a new species, is the complete ab-
sence of neural bones. Otherwise, the carapace is typical of other
South American representatives of the genus. The nuchal bone
is roughly pentagonal and slightly broader than long in its maxi-
mum dimensions; its postero-lateral borders are bowed outwards
1 The larger number represents the estimated total length of VF 11 77a;
its actual midline length as preserved is 60.1 centimeters.
1971
NEW VENEZUELAN FOSSIL PELOMEDUSID
slightly. There is no indentation at the midline of the anterior
margin. The eight pairs of pleurals meet in the midline. As in
all pelomedusids, there are eleven pairs of peripherals. The pygal
is trapezoidal and the suprapygal is roughly subtriangular. But-
tresses of the axial and inguinal notches are attached to the under-
sides of the first and fifth pleurals respectively. The iliac scars of
the pelvis are situated on the visceral surfaces of the seventh and
eighth pair of pleurals.
Of the three elements of the plastron, the bridge is the longest
while the anterior lobe is the shortest. The anterior plastral lobe
is U-shaped and does not extend beyond the anterior lip of the
carapace. The lateral margins of the posterior lobe are straight
rather than curved and are inclined medially so that the posterior
lobe becomes narrower toward the rear. The entoplastron is
diamond-shaped and has a slight U-shaped ridge with the open
end facing anteriorly on its visceral surface; this is presumably
for the attachment of neck muscles. At the midline junction of the
epiplastra there is a pronounced protuberance on the visceral side.
The mesoplastra are subrounded to hexagonal elements situated
^
^
Figure 3. The shape of the anal notch in three specimens of Podocnemis
veneziielensis (from left to right, VF 1177, VF 1173 and VF 1177a). The
arrow points toward the anterior end of the shell. The scale represents
a length of 10 centimeters.
8 BREVIORA No. 376
laterally at the base of the bridge. (These can best be observed
on the internal surface of VF 1174; see Plate V.) There is some
variation in the shape of the anal notch; in the smallest specimen,
VF 1 177, it is V-shaped, but in the two larger examples, VF 1 173
and VF 11 77a, it is broader at its base and thus tends to be more
U-shaped (Fig. 3). This structural difference may be due to sexual
dimorphism. If so, however, it is not possible to specify which
type represents the males and which the females in this species.
Normally, male turtles can be distinguished from females by the
characteristic depression on the posterior lobe of their plastrons,
but it is not possible to determine whether or not these existed in
the present sample because, in the course of fossilization, all the
plastra have been somewhat depressed inward along the midline.
Size alone is not a useful criterion for identifying the sexes in
Podocneinis. In all but one of the living South American species
of this genus mature females are invariably larger than males. The
reverse is true, however, for P. dumeriliana (F. Medem, personal
communication ) . Nor does the shape of the anal notch necessarily
provide a reliable method of sex determination. Although in at
least one species, P. lewyana, males can be identified solely on
the basis of this character, there are other species (e.g., P. vogli)
in which the two sexes appear to have similarly shaped anal
notches, while in still others (e.g., P. unifilis) the shape of the
anal notch varies considerably but evidently also randomly with
respect to sex.
Pelvic scars on the plastron appear to be disposed in essentially
the same positions as in the living South American species.
Although partial pelves have been preserved in several of the
specimens (VF 1173, 1176, 1177, and perhaps also 1177a), none
of these are well enough preserved to describe in detail.
The distal half of a right humerus (VF 1059) is all that is
known of the appendicular skeleton. Nothing serves to distinguish
this limb fragment from comparable portions of this same bone
in other species of Podocnemis.
Virtually no scute sulci can be detected in any of the specimens
available for study. This is disappointing because their arrange-
ment, particularly on the anterior plastral lobe, is sometimes of
taxonomic sianificance.
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 9
DISCUSSION
Taxonomic considerations. Within the suborder Pleurodira,
the presence or absence of neural bones has been accorded vary-
ing taxonomic significance. Those chelyid species that do possess
neurals usually have a variable number, and in certain species of
this family neurals evidently may or may not be present in diff"erent
individuals of the same population. Consequently, whether or
not a chelyid carapace includes some neurals, and if so, how many,
has never been considered a useful taxonomic character ^ Most
pelomedusid species, however, have a neural series that does not
deviate from a modal number (usually 6-8) by more than one
or, occasionally, two. Only two exceptions to this typical condition
are known within the family, both involving extinct taxa from the
Eocene of Tunisia-. Gajsachelys (de Stefano, 1903; Bergounioux,
1952; 1955; 1956), hke some chelyids, appears to have had a
rather variable number of irregularly shaped neurals. Eusarkia
(Bergounioux, 1952; 1956), described on the basis of a single
specimen, has no neurals and on the basis of this and several other
characters noted by Bergounioux we believe that it was probably
appropriate to propose a new genus. But in no case has the
absence of neurals alone been used as a taxonomic character,
either at the generic or the specific level, within the Pleurodira
(or among any other chelonians for that matter).
Why, then, have we described the Urumaco fossil pelomedusids
as a new species of Podocnemis? In view of their strong overall
resemblance to the living South American species of this genus
it seems inappropriate to propose a new genus on the basis of a
single character which, by itself, is not highly unusual nor of
particularly great taxonomic significance among other members
of the suborder. On the other hand, since the absence of neurals
is clearly a constant character within the Venezuelan sample, this
1 No adequate osteological descriptions of any living chelyid species
have ever been published, so that reliable data are not actually available
regarding the extent of intraspecific variation in the number of neurals.
-One of us (RCW) is preparing a discussion of the taxonomic status
of the Tunisian fossil turtles for publication elsewhere. In this paper
Euclastochelys (Bergounioux, 1955; 1956) is considered to be synonymous
with Gajsachelys.
10 BREVIORA No. 376
feature can hardly be regarded as an aberrant condition of no
taxonomic consequence. Thus, by a process of elimination, the
only alternative is to choose a procedure intermediate between
regarding the lack of neurals as of enormous taxonomic impor-
tance or as of none at all and describe the Venezuelan material
as a new species.
Ecological considerations. The small vertebrate fauna with
which Podocnemis veneziielensis is associated (Royo y Gomez,
1960: 509; Pascual and Diaz de Gamero, 1969: 370 and 374)
is not adequate for determining with any degree of certainty what
the probable habitat of this species might have been. The mam-
mals — a eumegamyine rodent and a toxodontid — were un-
doubtedly strictly terrestrial forms, while the crocodilians pre-
sumably spent most of their time in streams, lakes, or swamps.
The fish — sharks, sawfish, rays, catfish, and an unidentified
teleost — appear to be a mixture of marine and fresh water forms.
If all these fossils were collected from a single horizon, as the
scanty field evidence would suggest, then the stratum in which
they occur must represent an estuarine facies. On the basis of
present evidence, therefore, it is impossible to determine unequiv-
ocally whether P. veneziielensis was a marine or a fresh water
form. To be able to do so would be particularly interesting be-
cause, while all living pelomedusids are inhabitants of fresh waters,
in the past some were marine and others were fresh water forms
(Wood, MS). If P. veneziielensis were, in fact, marine, it would
be the last recorded pelomedusid so adapted.
No hving species of Podocnemis (or any other fresh water
turtle) are found in the Maracaibo basin, in which the type local-
ity of P. veneziielensis lies, although they are common to the
south and east of this enclave in Venezuela as well as to the west
of it in Colombia. Thus, P. veneziielensis occurs outside the pres-
ent range of the genus. Should P. veneziielensis eventually prove
to be a fresh water rather than a marine form, its extinction may
be explicable in terms of the Pleistocene climatic history of tropical
South America. The Maracaibo basin is ringed by mountains
except on its seaward side and hence is effectively isolated from
adjacent land areas. Conceivably, a period or periods of aridity
during the Pleistocene (and evidence for severe climatic fluctua-
tions in the tropics during this epoch is accumulating — cf . Van-
zolini and Williams, 1970: 94-103) may have eliminated P.
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 11
venezuelensis, a form presumably endemic to the basin, while the
surrounding mountain barrier prevented subsequent recolonization
by other species.
The fossil record of Podocnemis in South America. Few fossil
species of Podocnemis have been described from this continent
and several of these are known from such inadequate material
that it is questionable whether or not they should be referred to
the genus.
As previously noted, Simpson (1943) has described a partial
carapace and plastron from Venezuela as Podocnemis geolo-
gorum^. The single known specimen was recovered from fluvia-
tile beds of Miocene age. Without a doubt, this fossil represents
some kind of pleurodire because of the union of its pelvis with
both carapace and plastron. Whether this specimen actually rep-
resents a pelomedusid instead of a chelyid, however, is not entirely
clear. Although Simpson (1943: 57) commented "Es muy po-
sible la existencia en esta especie de un mesoplastron tipo Podoc-
nemis de buen tamafio," the presence or absence of this pair of
bones, the critical character for distinguishing members of one
pleurodiran family from the other, cannot be determined. Very
few potentially useful taxonomic characters can, in fact, be dis-
cerned. On the carapace, only two pleurals separate the last
neural from the suprapygal. There appears to be a deep indenta-
tion in the posterior edge of the pygal at the midline, and this,
together with similar but not quite so pronounced indentations
in the tenth and eleventh peripherals, gives the rear margin of
the carapace a serrated appearance. Whereas the last vertebral
was broader than long, the two preceding ones were longer than
broad. The anal notch of the plastron is rather deep and narrow,
and the lateral margins of the posterior lobe are sinuous. As a
consequence, the xiphiplastral tips are much more elongate than
in any other taxon yet described as a pelomedusid. Simpson
(1943: 61) considered the deep anal notch and serrations along
the posterior border of the carapace to be the species-specific
characters of P. geologorum, and indeed, these are quite distinctive
and indicate the validity of the species. Unfortunately, however,
1 When Simpson described the type of P. geologorum. it was catalogued
as AMNH 6781. It now belongs to the collections of the Museo de Ciencias
Naturales, Caracas, and bears the number MCN 915.
12 BREVIORA No. 376
on the basis of present evidence there is no reason to believe that
P. geologorum is really a species of Podocnemis or, for that mat-
ter, of any pelomedusid. It cannot yet be confidently allocated to
either of the two pleurodiran families to which it must belong,
the Chelyidae or the Pelomedusidae. Thus, until better material
of this taxon becomes available, P. geologorum should be listed
as Pleurodira incertae sedis.
Three species of Podocnemis — P. harrisi (Pacheco, 1913), P.
brasiliensis (Staesche, 1937)\ and P. elegans (Suarez, 1969) —
have been described from three widely separated localities within
the Bauri'i Formation of southern Brazil. These sediments repre-
sent terrestrial deposition, with fluvial and alluvial plain beds
predominating, and are probably of late Cretaceous (Senonian)
age (Oliveira, 1956: 53-54). The only specimens ever referred
to P. harrisi were a nearly complete right xiphiplastron and sev-
eral peripherals (Pacheco, 1913: 37, pi. 3 [figs. 6a-e], pi. 4
[fig. 6]). Ischial and pubic scars on the visceral surface of the
xiphiplastron clearly indicate that some kind of pleurodire is rep-
resented, but no other taxonomically useful evidence exists. It
is therefore impossible to determine whether P. harrisi is a chelyid
or a pelomedusid. Reference of this species to Podocnemis was
unjustifiable, as Schmidt noted long ago (1931: 253). Further-
more, since the type material is now apparently lost (Price, 1953:
10), "P. harrisi" must be regarded as a nomen vanum.
On the basis of some photographs of a partial plastron, a car-
apace fragment-, and three associated pleurals, Staesche (1937)
described P. brasiliensis. The xiphiplastra of this species do not
appear to dift'er in any appreciable way from those of P. harrisi,
and therefore Simpson (1943: 61) may well have been correct
in suggesting that P. brasiliensis is a synonym of P. harrisi,
although Staesche (1937: 302-303) noted that his material dif-
fered in that it represented a somewhat larger individual with a
difl'erent kind of sculpturing on the external surface of the shell,
factors which might or might not be of taxonomic significance.
1 Staesche (1944) is merely a translation of Staesche (1937) from
German into Portuguese.
-This carapace fragment, together with a previously undescribed ante-
rior lobe of a plastron from the same locality, was subsequently referred
to a new genus and species, Roxochelys wanderleyi, by Price (1953).
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 13
Small, laterally placed mesoplastra were present, even though no
longer preserved, so that P. brasiiiensis is clearly a pelomedusid.
An appropriate generic determination cannot at present be made,
however, because much of the anterior plastral lobe, so critical
for pelomedusid shell taxonomy, is missing. Thus, the specimen
can neither be certainly referred to Podocnemis nor, owing it its
imperfect preservation, can any species-specific characters be
established. Until better material is available, therefore, the plas-
tron and pleurals to which the name P. brasiiiensis now applies
should be designated as Pelomedusidae gen. et sp. indet. {"Podoc-
nemis brasiiiensis" Staesche).
Unlike the other two dubious "species" of Podocnemis from the
Baurii Formation, P. elegans is clearly valid and referable to this
genus. It is the only South American fossil pelomedusid yet
described for which associated shells and skulls have been re-
covered. Furthermore, it is the oldest representative of Podoc-
nemis known anywhere. It was described on the basis of two
specimens, an essentially complete shell and a well-preserved skull
belonging to a different individual. A detailed description of this
species will not be presented here since an account, based on addi-
tional new material as well as the original hypodigm, is being pre-
pared by one of us (RCW) for separate publication. Some of the
salient characters may be briefly noted, however. In most respects
the shell is typical of all South American species of Podocnemis,
but the shapes of the first two neurals are unique: instead of being
spindle-shaped, the first is hexagonal, with the postero-lateral sides
much shorter than the antero-lateral ones; and the second, rather
than being hexagonal, is subrectangular. A unique feature of the
skull is the total absence of triturating ridges on the palatal surface
of the upper jaw. All other species have from one to three triturat-
ing ridges, the exact number being characteristic of different spe-
cies. In addition, there does not seem to be an antero-posterior
1 Suarez (1969: 37) stated: "Designamos como tipo da nova especie a
carapaca e plastrao com craneo e diversos elementos esqueletais . . ."
The shell and skeletal elements belong to one individual and there is in
fact a badly crushed skull (which was not illustrated or discussed) asso-
ciated with them, but it is not the skull described by Suarez. This is an
isolated one from a much larger individual. Both specimens are in the
paleontological collections of the Faculdade de Filosofia. Ciencias e Letras
de Presidente Prudente: they bear no catalog numbers.
14 BREVIORA No. 376
forehead groove between the orbits. Of the other South American
species of Podocnemis, only P. dumeriliana lacks this groove.
Perhaps the most extraordinary aspect of the skull of P. elegans
is its modern appearance; archaic or ancestral features that one
might expect to find in such an ancient species are notably lacking.
Cattoi and Freiberg (1958) described Podocnemis argentinensis
from the Santa Barbara Formation^ in the Province of Jujuy,
Argentina. It is known from a large part of a plastron, lacking
the terminal portions of the anterior and posterior lobes as well
as much of the bridges, and most of the right epiplastron of a
second individual. The systematic position of this form is uncer-
tain. Laterally placed mesoplastra were definitely present, as
evidenced by the semicircular excavations on either side of the
plastron at the base of the bridge, so that argentinensis clearly
represents some kind of pelomedusid. Too little of the shell has
been preserved, however, to permit assignment to Podocnemis
with any degree of confidence. At a lower taxonomic level, the
characters used to define the species are in some cases questionable
and in others of little or no taxonomic significance. Cattoi and
Freiberg described the entoplastron as cordiform, but in their
figure and plate it appears to be quadrangular. Examination of
the specimen itself indicates that the entoplastron is slightly dam-
aged anteriorly and that it was probably diamond-shaped orig-
inally. Small, triangular gular scutes are characteristic of most
pelomedusids, but the relatively small intergular, which was prob-
ably pentagonal, is certainly reminiscent of the condition typical
of South American species of Podocnemis and the North American
Bothremys (Gaftney and Zangerl, 1968). The various scute pro-
portions cited by Cattoi and Freiberg in their diagnosis yield no
useful taxonomic information. Anastomosing vermiculations cover
the external plastral surface as in the majority of pelomedusid
1 Various ages have been assigned to this stratigraphic unit, formerly
referred to as the Margas Multicolores. Cattoi and Freiberg placed it in
the late Cretaceous, while Bardack (1961) considered it to be middle
Tertiary. The recent discovery of a mammal skull high in the formation
indicates a Paleocene or early Eocene age (R. Pascual, personal com-
munication). There is some uncertainty as to whether these beds are of
marine or terrestrial origin (Cattoi and Freiberg, 1958: 59).
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 15
genera'. Until better material becomes available, it will not be
possible to determine the systematic position of this turtle. In
the meantime, it must be referred to as Pelomedusidae gen. et sp.
indet. {'Podocnemis argentinensis" Cattoi and Freiberg).
The type and only specimen of Podocnemis bassleri (Williams,
1956) is a large, exceedingly well-preserved skull. It was collected
in eastern Peru from beds of the Contamana Group, which in-
cludes sediments believed to range in age from Eocene to possibly
at late as Pliocene. Williams, on the basis of information supplied
by Kummel, reported that the skull "came from the uppermost
part of the . . . group," which suggests that its age falls within
the latter part of the Tertiary. He further remarked that "The
fossil itself is so close to a Recent species as to tend to support
the latest date geologically permissible." The skull differs only
in minor details from that of the living P. expansa. It seems fairly
certain that P. bassleri was closely related, if not directly ancestral,
to this species.
Unfortunately, the evolutionary history of Podocnemis in South
America cannot be reconstructed on the basis of present informa-
tion. Only three fossil species — P. bassleri, P. elegans, and P.
venezuelensis — are of unquestionable validity. One of these,
P. bassleri, is clearly very closely related to P. expansa. The rela-
tionships of the other two extinct species to living South American
forms are uncertain, owing to their distinctive shell characters.
With the exception of P. lewyand'-, the living South American
species of Podocnemis are all strikingly similar in terms of shell
1 Cattoi and Freiberg's figure I shows the entoplastron as being nearly
encompassed by unusually large epiplastra. a condition unknown in any
other chelonians that we are aware of. However, their sketch does not
accurately represent the positions of the sutures between the epiplastra
and hyoplastra; these are, in fact, disposed in typical pelomedusid fashion,
extending outward from the lateral apices of the entoplastron.
-Through the courtesy of Professor F. Medem, one of us (RCW) has
been able to examine a series of six P. lewyana shells in the collections
of the Instituto Roberto Franco at Villavicencio, Colombia. None of these
has a suprapygal bone on the carapace; instead, each of the last (eighth)
pleurals is subtriangular, not trapezia! as is the case in other species of the
genus, and these pleurals are in continuous contact along the midline from
the posterior end of the seventh pair of pleurals to the pygal.
16 BREVIORA No. 376
morphology. Only small structural details characteristic of each
taxon permit differentiation among them on the basis of shells
alone. P. venezuelensis stands markedly apart from all other spe-
cies of the genus in its total lack of neurals, and certainly could
not have given rise to any of the living forms. Nothing remotely
resembling the shapes of the first two neurals in P. elegans is en-
countered elsewhere in the genus. So conservative in structure is
this part of the shell in all other species (except, of course, for
P. venezuelensis ) , and so radically different is it in P. elegans, that
this species could hardly have been ancestral to any or all of the
later species known from South America. Thus, neither P. elegans
nor P. venezuelensis has any obvious relationship to living species
of the genus or to each other. A much better fossil record for
Podocnemis will be necessary before a meaningful picture of its
evolutionary history in South America can be formulated.
ACKNOWLEDGMENTS
We are particularly grateful to Senora Frances Charlton de
Rivero, retired professor of paleontology at the Escuela de Geo-
logia y Minas in Caracas, not only for her gracious hospitality but
also for having provided laboratory facilities while the material
here described was being studied. Both authors have examined
the type specimen of "Podocnemis'' geologorum and one of us
(RCW) has also been able to examine the types of "P. argen-
tinensis," P. bassleri, and "P. brasiliensis." We would like to thank
the curators of the various institutions at which these fossils are
housed for permission to study them. Through the kindness of
Professor Jose Martin Suarez, RCW has not only been able to
study the type material of P. elegans but also to visit the locality
from which it was recovered. Without a generous grant to RCW
from the National Geographic Society, none of this work would
have been possible. We are much obliged to Professors F. Medem,
B. Patterson, P. E. Vanzolini, and E. E. Williams, and to Dr. M.
Freiberg for their comments and discussions on various aspects
of this manuscript.
1971 NEW VENEZUELAN FOSSIL PELOMEDUSID 17
LITERATURE CITED
Bardack, D. 1961. New Tertiary teleosts from Argentina. American
Mils. Novitates. No. 2041: 1-27.
Bergounioux, F. M. 1952. Les cheloniens fossiles de Gafsa. Appendix
to: Arambourg, C, and J. Signeux. Les vertebres fossiles des gise-
ments de phosphates ( Maroc-Algerie-Tunisie). Notes et Memoires
No. 92, Service Geologiqiie de Maroc: 377-396, pis. 45-46.
. 1955. La famille des Eusarkiides. Comptes Rendus de
I'Acad. Sci.. 240: 1455-1457.
1956. Les reptiles fossiles des depots phosphates sud
tunisiens. Annales des Mines et de la Geologie (Tunis), No. 15:
1-105, pis. 1-17.
Cattoi, N., and M. a. Freiberg. 1958. Una nueva especie de "Podoc-
nemis" del cretaceo argentino. Physis. 21(60): 58-67.
Dacque, E. 1912. Die fossilen Schildkroten Aegyptens. Geol. Palaont.
Abhandl., N. F., 10(4): 275-337, pis. 36-37.^
Gaffney, E. S., and R. Zangerl. 1968. A revision of the chelonian
genus Bothremys (Pleurodira, Pelomedusidae). Fieldiana. Geol.,
16(7): 193-239.
Oliveira, a. I. DE. 1956. Brazil. //; W. F. Jenks (ed.). Handbook of
South American Geology. Geol. Soc. Amer. Mem. 65: 1-378.
Pacheco, J. A. 1913. Notas sobre a geologia do valle do Rio Grande, a
partir da foz do Rio Pardo ate a sua confluencia com o Rio Paranahyba.
/// J. dos Dourados (ed.). Explora^ao do Rio Grande e de seus afflu-
entes. Commisao Geographica e Geologica do Estado de Sao Paulo:
33-38.
Pascual, R.. and M. L. Diaz de Gamero. 1969. Sobre la presencia del
genero Eiimegamys (Rodentia, Caviomorpha) en la formacion
Urumaco del Estado Falcon (Venezuela). Su signification cronologica.
Bol. Informativo, Assoc. Venezolana de Geol.. Min. Pet.. 12(10):
369-387.
Price, L. I. 1953. Os quelonios de forma<:ao Baurii, Cretaceo terrestre
do Brasii meridional. Dept. Nac. Prod. Min., Div. Geol. Min., Bol.
147: 1-34.
RoYO Y Gomez, J. 1960. Les vertebrados de la formacion Urumaco,
estado Falcon. Mem. Ill Congr. Geol. Venezolano, 2: 506-510.
Schmidt, K. P. 1931. A fossil turtle from Peru. Field Mus. Nat. Hist.
Publ. 299, Geol. Ser., 4(8): 251-254.
1940. A new turtle of the genus Podocneniis from the
Cretaceous of Arkansas. Geol. Ser. Field Mus. Nat. Hist.. 8( 1): I-I2.
18 BREVIORA No. 376
Sill, W. D. 1970. Nota preliminar sobre un nuevo gavial del Plioceno
de Venezuela y una discusion de los gaviales sudamericanos.
Ameghiniana, 7(2): 151-159.
Simpson, G. G. 1943. Una tortuga del Terciario de Venezuela. Rev
Ministerio de Fomento, Caracas, Venezuela, ano 5, nos. 51-52,
April -Sept. 1943: 53-64.
Staesche, K. 1937. Podocneinis hrasiliensis n. sp. aus der Oberen Kreide
Brasiliens. Neues Jahrb. Min., Geol. Palaont., 77: 291-309.
. 1944. Uma tartaruga do cretaceo superior do Brasil. Dep.
Nac. Prod. Min., Div. Geol. Min., Bol. 114: 1-24, pis. 16-20.
Stefano, G. de. 1903. Nuovi rettili degli strati a fosfato della Tunisia.
Bol. Soc. Geol. Ital., 22: 51-80.
Suarez, J. M. 1969. Urn quelonio da formagao Bauru. Dept. Geografia,
Fac. de Filos., Cien. Letras Pres. Prudente, No. 2: 35-54.
SwiNTON, W. E. 1928. Note on the fossil reptilia collected by Mr. Don-
ald Stewart in Venezuela. Quart. Jour. Geol. Soc, 84, Pt. 3: 583.
Van Frank. R. 1957. A fossil collection from northern Venezuela.
1. Toxodontidae (Mammalia, Notoungulata). American Mus. Novi-
tates. No. 1850: 1-38.
Vanzolini, p. E., and E. E. Williams. 1970. South American anoles:
the geographic differentiation and evolution of the Anolis chrysolepis
species group (Sauria, Iguanidae). Arq. Zool.. S. Paulo, 19(1-2):
1-124.
Williams, E. E. 1956. Podocneinis bassleri, a new species of pelomedusid
turtle from the late Tertiary of Peru. American Mus. Novitates, No.
1782: 1-10.
Zangerl, R. 1948. The vertebrate fauna of the Selma Formation of
Alabama, part 11. The pleurodiran turtles. Fieldiana: Geol. Mem.
3(2): 19-56.
197
NF.W VENEZUELAN FOSSIL PELOMEDUSID
19
15 cm
Plate L Dorsal view of the type carapace (VF 1176) of Podocnemis
veneziielensis.
20
BREVIORA
No. 376
15 cm
Plate II. View of the visceral surface of the type carapace (VF 1176)
of Podocnemis venezuelensis. Note the absence of neural bones.
1971
NEW VENEZUELAN FOSSIL PELOMEDUSID
21
-^
^•^^.
V:^''
i
t-r '■
15 cm
Plate ill. View of the visceral surface of the type plastron of Pocloc-
neinis veneziielensis (VF 1173). showing the disposition of the pelvic
attachments to it.
22
BREVIORA
No. 376
■^- '^■■<%5
15 cm
Plate IV. Ventral view of the type shell of Potlocneiuis vcnezuelensis,
with the plastron (VF 1173) positioned correctly in relation to the carapace
(VF 1176).
1971
NEW VENEZUELAN FOSSIL PELOMEDUSID
23
Plate V. The visceral surface of VF 1174. a partial plastron of Podoc-
nemis venezneleii.sis, showing clearly the outlines of the laterally placed
mesoplastra.
BREVIORA
MmseiLim of Comparative Zoology
CAMBRrocE, Mass. 15 June. 1971 Number 377
THE CHANARES (ARGENTINA) TRIASSIC REPTILE FAUNA
IX. THE CHANARES FORMATION
Alfred Sherwood Romer
Abstract. The term "Chanares Formation" should be retained for the
fossihferous beds so named by Romer and Jensen; the type section of the
"Ischichuca Formation" is homologous with part of the overlying Los
Rastros Formation.
With the discovery of a rich Triassic reptilian fauna in the
Chafiares-Gualo region of La Rioja Province, Mr. James Jensen
and I set about the task of determining the stratigraphic sequence
of the area (Romer and Jensen, 1966). It was soon apparent that,
despite complex faulting, a series of formations with clear-cut
boundaries could be distinguished, and in the Arroyo del Agua
Escondida the entire local series could be seen in proper sequence.
The area is part of a basin of late Paleozoic and early Mesozoic
deposition lying in western La Rioja Province and an adjacent
portion of San Juan, extending roughly from the western flanks
of the Sanogasta Range on the east to the Rio Bermejo on the
west, and from the region of Villa Union south to the northern
end of the Valle Fertil range. The center of this area is the flat
Campo de Talampaya, and I shall term this cuenca the Talampaya
Basin. Little attention had ever been given to the geology of our
area of interest in the eastern part of the basin, but considerable
work had been done in the western part. It was obvious that the
thickness and nature of the sediments varied greatly from one
basin area to another, and the situation was further complicated by
the fact that there had been much volcanic activity. As far as
possible we utilized formation names already in the literature;
\vhcn no similarities to named formations in other areas in the
hasin were discernible, new names were given. The formations
named by us are shown in the right hand column of the table. The
2 BREVIORA No. 377
three formations latest in time — Los Colorados, Ischigualasto^
and Los Rastros — are comparable to those so named to the west,
across the Campo de Talampaya, although much thinner in our
area in each instance. Below the normal beds of the Los Rastros,
strongly carbonaceous in nature, are some 70 meters of evenly
bedded volcanic ash sediments in which fossil reptiles are abun-
dant. Despite the fact that these strata are quite conformable
with the overlying Los Rastros deposits, they are, as layers of
white to bluish white ash, quite distinctive in character and, since
they are the bearers of our fauna, we ventured to separate them
from the Los Rastros